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AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3033, 68 pp., 44 figures February 24, 1992

Phylogenetic Analysis and Taxonomy of the Tropidurus Group of Lizards (Iguania: Tropiduridae)

DARREL R. FROST"2

ABSTRACT Phylogenetic relationships among 44 species of groups) are parts of a single monophyletic group. the South American Tropidurus group of lizards Excepting Uranoscodon, all species of the Tropi- are analyzed using standard cladistic techniques. durus group east of the Andes are part of a single Seventy-seven transformation series of osteology, monophyletic group. Microlophus is resurrected squamation, color, and hemipenes are polarized for former species ofTropidurus west ofthe Andes, (when possible) using as first and second outgroups excepting T. koepckeorum, which is placed in a the Stenocercus group and Leiocephalus. Thirty- monotypic genus Plesiomicrolophus, in polytomy six equally parsimonious trees (length 169, ci = with Microlophus and Tropidurus. Tropidurus is 0.568) are discovered, of which one is also the redefined to include as synonyms Plica, Strobi- strict and Adams consensus tree of the other 35. lurus, Uracentron, and Tapinurus. Two new tribes Tropidurus is demonstrated to be paraphyletic with are diagnosed, Tropidurini, equivalent to the respect both to Tapinurus and to a monophyletic Tropidurus group, and Stenocercini, equivalent to group composed of Plica, Strobilurus, and Ura- the former Stenocercus group ("Ophryoessoides," centron. With the exception of T. koepckeorum, "Stenocercus," and Proctotretus). Within Steno- all species of Tropidurus west of the Andes (the cercini, Proctotretus and Ophryoessoides are syn- former T. occipitalis and T. peruvianus species onymized with Stenocercus.

INTRODUCTION Tropiduridae is a medium-sized family of three subfamilies: Liolaeminae (Liolaemus, Neotropical iguanian lizards composed of Ctenoblepharys, Phymaturus), Leiocephali-

1 Assistant Curator, Department of Herpetology and Ichthyology, American Museum of Natural History. 2This paper developed from part of a dissertation completed at the Museum of Natural History, University of Kansas.

ACKNOWLEDGMENTS For loan of and/or access to specimens I thank: Charles W. Myers and Richard G. Zweifel, American Museum of Natural His- tory (AMNH) (prior to my joining the staff at the AMNH); E. Nicholas Arnold, The Natural History Museum, London (BMNH); Robert C. Drewes, Jacques Gauthier, Alan Leviton, and Jens Vindum, California Acad- emy of Sciences (CAS); Raymond Laurent, Fundacion Miguel Lillo (FML); William E. Duellman, Joseph T. Collins, and John E. Simmons, Museum ofNatural History, Uni- versity of Kansas (KU); Robert L. Bezy and John W. Wright, Natural History Museum of Los Angeles County (LACM); Douglas A. Rossman, Museum ofNatural Science, Lou- isiana State University (LSUMZ); Mark No- rell, American Museum of Natural History (MAN private osteology collection); Ernest E. Williams and Jose Rosado, Museum of Comparative Zoology, Harvard University (MCZ); Teresa C. de Avila Pires, Museu Pa- Fig. 1. Distribution of the Tropidurus group. raense Emilio Goeldi (MPEG); Stephen D. Busack, David Good, Harry W. Greene, and David Wake, Museum of Vertebrate Zoolo- nae (Leiocephalus), and Tropidurinae. Tropi- gy, University of California (MVZ); Richard durinae is composed of two monophyletic Etheridge, San Diego State University (REE taxa, the Stenocercus group ("Ophryoes- private osteology collection and SDSU); soides,"3 "Stenocercus," Proctotretus) and the Gregory K. Pregill, San Diego Natural His- Tropidurus group (fig. 1) (Tropidurus, Tapi- tory Museum (SDSNH); James R. Dixon, nurus, Uranoscodon, Uracentron, Strobilu- Texas A&M University (TCWC); Arnold G. rus, and Plica) (Etheridge and de Queiroz, Kluge and Greg Schneider, Museum of Zo- 1988; Frost and Etheridge, 1989). Etheridge ology, University of Michigan (UMMZ); and de Queiroz (1988), Frost and Etheridge Norman J. Scott, University of New Mexico (1989), and Pregill (in press) have substan- (UNM); Ronald I. Crombie, Roy W. Mc- tiated the sister-taxon relationship of these Diarmid, Robert Reynolds, and George R. two groups, and the relationship ofthe Tropi- Zug, National Museum of Natural History durinae with the Leiocephalinae (fig. 2). The (USNM); and Jonathan A. Campbell, Uni- purpose of this study is to formulate a hy- versity of Texas at Arlington (UTA). pothesis based on all available evidence, Richard Etheridge, Gregory K. Pregill, and compare the results with the current hypoth- John W. Wright have been of considerable esis ofintragroup relationships (Etheridge and help through their discussions and com- de Queiroz, 1988), and provide a taxonomy ments, although this should not be construed logically consistent (Hull, 1964; Wiley, 1981) necessarily as acceptance of conclusions in with recovered history. this manuscript. Besides these, several others have provided helpful criticism ofthis manu- at stages of its development: 3Quotation marks surround names of supraspecific script various taxa that are not demonstrably monophyletic (Wiley, Maureen A. Donnelly, William E. Duellman, 1979; Kluge, 1989a). For purposes of clarity, all taxa in Arnold G. Kluge, Mathias Lang, Roy W. the Tropidurus group will be considered monophyletic McDiarmid, Tom Titus, Linda Trueb, and unless demonstrated to be otherwise. E. 0. Wiley. I also thank John Cadle for let- 1992 FROST: TROPIDURUS GROUP OF LIZARDS 3 ting me read his Stenocercus manuscript; his tural cluster: Lacerta plica (now Plica plica), comments therein regarding some overstate- L. umbra (now Plica umbra), and L. super- ments in my dissertation have helped me to ciliosa (now Uranoscodon superciliosus). refine this manuscript. Anne Musser execut- Laurenti (1768) provided a junior syn- ed figure 18. Permission to use photographs onym of L. plica in his Iguana chalcidica. was granted by: David C. Cannatella, Martha Latreille (1801) placed L. umbra and L. su- L. Crump, James R. Dixon, William E. perciliosa in Iguana, and L. plica in Stellio. Duellman, Roy W. McDiarmid, Charles W. In the following year, Daudin (1802) consid- Myers, Laurie J. Vitt, and Richard G. Zwei- ered all three species to be in the genus Aga- fel. ma, along with most smaller scansorial non- chameleon iguanians. Merrem (1820) also HISTORICAL REVIEW included all three species under Agama, but supplied a substitute name, Agama tigrina, Prior to the publication of Etheridge's in- for Iguana superciliosa Latreille, 1801 (= L. formal tree ofrelationships within the former superciliosa Linnaeus, 1758). Oppel (1811) "Iguanidae" (Etheridge in Paull et al., 1976), transferred Daudin's (1802) species of Aga- questions of phylogeny in the Tropidurus ma and Iguana into Lophyrus Dumeril group (Uranoscodon, Tropidurus, Plica, Ta- (1806). pinurus, Strobilurus, and Uracentron) were Kaup (1825) erected the name Uranosco- imbedded in larger discussions of relation- don (unjustifiably emended to Uraniscodon ships within the former "Iguanidae" (i.e., by Boie, 1825), based on L. superciliosa, but Crotaphytidae, Corytophanidae, Hoplocer- also including the Linnaean species L. plica cidae, Iguanidae, Phrynosomatidae, Poly- and L. umbra. Boie (1825) suggested that chridae, Tropiduridae) and former tropidu- Agama catenata Wied-Neuwied (1821) and rine iguanids (= Tropiduridae) (Etheridge, Agama picta Wied-Neuwied (1825) (both 1964, 1966, 1967). species currently in Enyalius [Polychridae]) Boulenger (1885) listed the genera of the should be included in a genus with L. super- Tropidurus group nonalphabetically, thereby ciliosa. He noted that his name Ophryessa, implying some concept ofrelationship, but it or Uraniscodon, could be used. However, he was not until Etheridge and de Queiroz (1988) excluded plica and umbra from this genus. published their systematic review of "Igua- Spix (1825) named Lophyrus ochrocollaris nidae" (sensu lato) that the Tropidurus group, (ajunior ofLacerta umbra, considered a sub- within tropidurine iguanids, had been diag- species ofP. umbra by Etheridge, 1970a), Lo- nosed sufficiently well to permit detailed dis- phyrus panthera (a junior synonym of La- cussion. Subsequently, Frost and Etheridge certa plica), and Lophyrus xiphosurus and (1989) partitioned the likely paraphyletic Lophyrus aureonitens (both junior synonyms "Iguanidae" and recognized these genera of Lacerta superciliosa) from Brazil. Subse- (listed above) as parts ofone ofits constituent quently, Kaup (1826) synonymized Lophyrus groups, Tropiduridae. xiphosurus Spix and L. aureonitens Spix with A discussion of the taxonomic history of L. superciliosa Linnaeus. He also provided a the genera and species ofthe Tropidurus group more detailed characterization of the genus is hampered by its complexity and historical (as Uraniscodon) and included, in addition confusion. Because of this, I will discuss this to superciliosa, picta, and umbra, a number history in convenient nomenclatural clusters: ofnames currently referable to Enyalius and Uranoscodon and Plica; Uracentron and Tropidurus hispidus (Spix). Boie (1825), deal- Strobilurus; and Tapinurus and Tropidurus. ing with the same species, placed in Ophryes- sa the species superciliosa, ochrocollaris (= Uranoscodon and Plica umbra), panthera (= plica), aureonitens (= Etheridge (1970a) provided an extensive superciliosa), catenata (= Enyalius catena- review ofthe taxonomic history ofPlica; with tus), and margariticeus (= Enyalius pictus). minor changes that review is paraphrased Fitzinger (1826) recognized a new genus, here. Linnaeus (1758) named all of the cur- Ecphymotes (type species subsequently des- rently named species within this nomencla- ignated by Fitzinger, 1843, as acutirostris), 4 AMERICAN MUSEUM NOVITATES NO. 3033

for plica, undulatus (= Anisolepis undulatus plica. In the same work they erected a new [Polychridae]), pictus (= Enyalius pictus genus, Uperanodon, to include part of Plica [Polychridae]), and acutirostris (= Polychrus Gray and part of Hypsibatus Wagler: ochro- acutirostris [Polychridae]), but he retained collare (= P. umbra) and pictum (= Enyalius Boie's Ophryessa, in which he included su- pictus). Ophryoessa (an unjustified emenda- perciliosa, catenatus (= Enyalius catenatus), tion of Ophryessa) was used by these authors margaritaceus (= Enyalius catenatus), and to accommodate the single species supercilio- umbra. Kaup (1827) responded by recom- sa. In the same year, Gravenhorst (1837) fol- mending that Uraniscodon be partitioned into lowed Wiegmann (1835) and used Hypselo- three subgenera: Uraniscodon forAgama his- pus (an unjustified emendation) as the generic pida Spix (= Tropidurus hispidus); Pneustes name for plica. for picta, umbra, and plica; and Ophryessa Fitzinger (1843) treated Enyalius (for cate- for superciliosa, catenata, and margaritaceus. natus and margaritaceus), Hypsibatus (for The number of subgenera recommended (3) umbra), Uperanodon (for pictus), Dryophilus is probably significant, because Kaup was pre- (for bilineatus), and Ophryoessa (for super- evolutionary in his views, believing that 3s ciliosa) as subgenera within Hypsibatus. He and 5s had a deep natural significance (Mer- also erected a new genus, Ptychosaurus. Be- tens, 1973). cause Fitzinger recognized two synonyms of Wagler (1830) erected the genus Hypsiba- plica as distinct species, he erected different tus for plica, umbra, and picta. Subsequently, subgenera within Ptychosaurus for each: Pty- Wiegmann (1835) replaced Hypsibatus with chosaurus (for Hypsibatuspunctatus Dumeril Hypselophus, thinking that Hypsibatus was and Bibron) and Ptychopleura (for L. plica). preoccupied by Hypsibates Nitsch. Under the A third subgenus of Ptychosaurus, Tritopis, current Code ofNomenclature (1985) no such was erected for Tropidogaster blainvillii Du- preoccupation exists. However, Etheridge meril and Bibron (1837) (= Chalarodon mad- (1970a) considered, by reason ofArticle 23b agascariensis, fide Etheridge, 1969a). In 1864, ofthe 1961 International Code ofZoological Fitzinger regarded his subgenus Ptychopleura Nomenclature, that Hypsibatus was preclud- (ofPtychosaurus) as a genus distinct from the ed for competing in synonymy by reason of nominate subgenus, even though they had the its long lack of use. Contrary to a comment same biological type species. made by Frost and Etheridge (1989), this sup- Gray (1845) referred umbra and picta to pression is continued under the 1985 Code Uraniscodon, but referred umbra (under the by Article 79(c)iii which states that actions misidentification of plica) and punctata (= taken under 23b of the 1961 Code are to be plica) to Plica. Following Fitzinger (1843), he upheld unless acted upon by the Commis- retained superciliosa in Ophryoessa (an un- sion. justified emendation of Ophryessa). Gray (1827) described Lophyrus aga- Boulenger (1885) was the first to group to- moides, which he later (1831) considered to gether exclusivelyplica and umbra -albeit in be a synonym of Lacerta plica. Plica was Uraniscodon. Ophryoessa was considered to erected by Gray (1831) as a subgenus of be a monotypic genus for superciliosa. Stejne- Ophyessa (an unjustified emendation of ger (1901) noted the incongruity of the ar- Ophryessa Boie) to include brasiliensis (= En- rangement; Uranoscodon Kaup was based on yalius catenatus [Polychridae]), picta (= Plica superciliosa, even though Kaup included both umbra, in this case, according to Etheridge, umbra and plica. Boie's (1825) Uraniscodon 1970a), and plica. Gray erected a subgenus, included only superciliosa of the originally Xiphura, of Ophyessa, for superciliosa, mar- included species; Stejneger (1901) regarded garitaceus (= Enyaliuspictus), and rhombifer this as tantamount to fixation of a type spe- (= Enyalius catenatus). Gray's confusion be- cies. Burt and Burt (1931) followed Stejneger tween Agama picta Wied-Neuwied (= Eny- (1901) in the use of the name Uranoscodon alius pictus) and Plica plica and P. umbra superciliosa. Etheridge (1970a) formally des- was continued by Schinz (1833). ignated L. superciliosa Linnaeus as the type Dumeril and Bibron (1837) employed species of Uranoscodon, and placed three Hypsibatus Wagler for H. agamoides and the names in the synonymy ofPlica umbra: Tro- new H. punctatus, both synonyms of Plica pidurus unicarinatus Werner (1899), T. hol- 1 992 FROST: TROPIDURUS GROUP OF LIZARDS 5 otropis Boulenger (1912), and Plica tubercu- along with other spiny-tailed species, to Ho- latum Andersson (1918). He also excluded plurus. one species, Plica stejnegeri Burt and Burt O'Shaughnessy (1881) correctly placed Do- (1930) (= Tropidurus spinulosus), from Plica. ryphorus flaviceps Guichenot in Uracentron Etheridge's (1 970a) revision of Plica was the (as Uranocentrum). Boulenger (1885) recog- first to characterize the genus adequately. nized three species of Uracentron: U. azure- Most recently, Plica lumaria was discovered um (Linnaeus, 1758), U. flaviceps (Guiche- and named by Donnelly and Myers (1991) not, 1855), and U. castor (Cope, 1870). Also, and (as Plica nigra, a junior synonym) by he recognized that Doryphorus spinosus is a Miigdefrau (1991). This species, phenotypi- junior synonym of Strobilurus torquatus. cally similar but also plesiomorphic in some Burt and Burt (1933), following Fitzinger respects ofsquamation to Plica plica, is from (1843) and Tschudi (1845), confused Ura- Cerro Guaiquinima in southern Venezuela. centron and Phymaturus, and mistakenly included palluma in Uracentron. They also included Urocentrum meyeri Werner (1900), Uracentron and Strobilurus which subsequently was demonstrated to be Linnaeus (1758) named Lacerta azurea, a member of Stenocercus (Etheridge, 1968). mistakenly thought to be from Africa. Sub- Etheridge (1968) reviewed both Strobilurus sequently, Latreille (1802) named Stellio and Uracentron. He maintained the mono- brevicaudatus, ajunior synonym ofL. azurea. typic status ofStrobilurus and recognized four He placed this species and L. azurea in Stel- species in Uracentron: azureum, guentheri, lio, along with other spiny-tailed lizards. This werneri, andflaviceps. Greene (1977) reeval- arrangement was followed by Daudin (1802) uated the status of the species of Uracentron and Fitzinger (1826). Merrem (1820) placed and regarded guentheri and werneri as sub- azurea in Uromastyx (another genus ofspiny- species of azureum. tailed lizard), and supplied a replacement name, Uromastyx caeruleus. Kaup (1826) introduced the name Uracen- Tropidurus and Tapinurus tron for azurea (and caerulea). Wagler (1830) Unlike those in the other genera within the emended the name to Urocentron, which en- Tropidurus group, most of the species in joyed common usage prior to 1968 (e.g., Fitz- Tropidurus and Tapinurus have become inger, 1843; Mertens, 1925; Dunn, 1944; known only relatively recently. Wied-Neu- Valdivieso and Tamsitt, 1963; Peters, 1967). wied (1820) described Stellio torquatus from Other emendations of Uracentron were Ura- Brazil; subsequently, he (1825) erected a new nocentron (Gray, 1831, 1845), Uranocen- genus, Tropidurus, for this species. Lichten- trum (O'Shaughnessy, 1881), and Urocen- stein (1822) named torquatus (fide Ro- trum (Boulenger, 1894; Werner, 1900). Cuvier drigues, 1987) as Agama operculata, and (1829) supplied Doryphorus as a substitute Raddi (1822) named it as Agama brasiliensis. name, and it enjoyed some popularity (Schinz, Spix (1825) named several new species from 1835; Dumeril and Bibron, 1837; Guichenot, Brazil: Agama hispida (now Tropidurus his- 1855; Dumeril, 1856; Cope, 1870). Gray pidus), A. nigrocollaris (a synonym ofA. his- (1831) placed Uracentron (as Uranocentron) pida, fide Peters, 1877), A. cyclurus (a syn- as a subgenus of Ophryessa, along with Plica. onym of A. hispida, fide Peters, 1877), A. Wiegmann (1 834b) named a new genus and semitaeniatus (now Tapinurus semitaenia- species ofspiny-tailed lizard, Strobilurus tor- tus), and A. tuberculata (a synonym of Tropi- quatus, from Brazil. Guichenot (1855) in- durus torquatus, fide Rodrigues, 1987). Kaup advertently renamed this species as Dorypho- (1826) mistakenly transferred A. hispida into rus spinosus, but in the same paper he named Uraniscodon with species currently allocated a second species, Doryphorus flaviceps, that to Uranoscodon and Plica. is currently in Uracentron. Fitzinger (1843) Fitzinger (1826) recognized Tropidurus placed Strobilurus as a subgenus of Steiron- Wied-Neuwied (1820), but he included otus, which also included a few species cur- schreibersi (nomen nudum = Pristinotus rently allocated to Leiocephalus and Steno- schreibersi Gravenhorst, 1837 = Leiocepha- cercus. Schlegel (1858) transferred azurea, lus schreibersi), along with torquatus. 6 AMERICAN MUSEUM NOVITATES NO. 3033

Wagler (1830) erected Platynotus for Aga- tigris. In the same year, Gray (1845) replaced ma semitaeniata Spix, and recognized an en- Tropidurus with a new name, Taraguira, and larged Tropidurus that contained torquatus added two new species names: smithii (= his- (including tuberculata and hispida as syn- pidus) and darwini (= torquatus). He recog- onyms), nigrocollaris (with cyclurus consid- nized Microlophus for peruvianus and trans- ered a synonym), and a few other species cur- ferred grayii to Leiocephalus in the subgenus rently allocated to the generalized scansorial Holotrophis. genera Sceloporus and Oplurus. This arrange- Gray (1845) named two junior synonyms, ment was followed by Gray (1831), who re- Taraguira smithii (= Tropidurus hispidus, fide tained A. hispida Spix as a distinct species, Boulenger, 1885) and Taraguira darwinii (= but reallocated semitaeniatus to Tropidurus. T. torquatus, fide Boulenger, 1885). Berthold Lesson (1831) named Stellioperuvianus and (1859) named another junior synonym of Lophyrus araucanus (both = peruvianus, fide Tropidurus hispidus, Proctotretus toelsneri, Ortiz-Zapata, 1980a), the first species cur- and in 1861, two more were named: Trachy- rently allocated to Tropidurus to be named cyclus superciliaris Gunther (1861) and Trop- from west of the Andes. Wiegmann (1834a) idurus macrolepis Reinhardt and Liitken named two more species from west of the (1861). Reinhardt and Liitken (1861) also Andes, Tropidurus heterolepis and T. micro- named a currently recognized species, T. hy- lophus (= peruvianus), and for the first time gomi, from Brazil, as well as Tropidurus mac- associated species from both sides ofthe An- rolepis (= T. hispidus, fide Rodrigues, 1987). des under Tropidurus. Wiegmann (1834b) Cope (1862) named Microlophus spinulosus continued to recognize Platynotus semitaeni- (now Tropidurus spinulosus), from Paraguay. atus and maintained Wagler's (1830) concept With remarkable insight, Peters (1871) re- of Tropidurus, only adding the two new spe- turned grayii to Tropidurus and named two cies and some species currently referable to new species, bivittata (from the Galapagos) Liolaemus. and occipitalis (from western Ecuador). He Dumeril and Bibron (1837) erected Micro- also conceived of Tropidurus as composed of lophus for their new species M. /essonii; they four subgenera: Craniopeltis (grayii and bivit- regarded all other species from west of the tata), Laemopristis (occipitalis), Microlophus Andes (i.e., peruvianus, microlophus, heter- (microlophus and heterolepis), and Tropidu- olepis), as synonyms. For Tropidurus torqua- rus (torquatus and macrolepis). tus they suggested Ecphymotes (not of Fitz- Bocourt (1874) named Aneuoporus occip- inger, 1826) as a replacement generic name. italis, which is identical to Laemopristis oc- Surprisingly, I have been unable to find any cipitalis Peters (1871). mention ofA. semitaeniata Spix in this clas- Steindachner (1876) described Tropidurus sic work. (Craniopeltis) pacificus pacificus and T. (C.) Bell (1843) named the first of the Gala- p. habelii from the Galapagos, following Pe- pagos Tropidurus from material collected by ters' (1871) subgeneric arrangement. In the Darwin as Leiocephalus grayii. Fitzinger same year, Cope (1876) placed occipitalis into (1843) erected Steirolepis as a rough equiv- Craniope/tis, thereby synonymizing Laemo- alent of Microlophus Dumeril and Bibron pristis Peters. In the same paper, Cope named (1837), including microlophus, heterolepis, Microlophus inguinalis (= peruvianus) and andperuviana, but also including semitaenia- transferred heterolepis to Microlophus. ta, presumably because semitaeniatus shares Gunther (1877), staying with Bell's (1843) small scales with these other taxa. He re- earlierjudgment, returned pacificus and gray- tained Tropidurus for torquatus (including ii (including bivittata) to Leiocephalus. In the hispida and tuberculata as synonyms) and same year, Peters (1877), regarded Platynotus microlepidotus (probably a synonym of tor- as a subgenus of Tropidurus. quatus; including nigrocollaris and cyclurus). O'Shaughnessy (1879), following Gunther Tschudi (1845) followed Fitzinger's (1843) (1877), named Liocephalus (Craniope/tis) use ofSteirolepis and added four new species variegatus, a junior synonym ofMicrolophus names from Peru: xanthostigma (= peruvi- spinulosus Cope, 1862. Boettger (1885) fol- anus), quadrivittata (type locality in a region lowed Peters (1871) and regarded Microlo- transferred to Chile in 1878), thoracica, and phus as a subgenus of Tropidurus. 1 992 FROST: TROPIDURUS GROUP OF LIZARDS 7

In the Catalogue ofLizards in the British Tropidurus. They recognized seven species in Museum (NaturalHistory), Boulenger (1885) the Galapagos: pacificus, duncanensis, ha- considered all ofthe generic names based on belii, bivittatus, delanonis, grayii, and albe- species within the nomenclatural cluster marlensis (with two subspecies, albemarlen- composed of current Tropidurus and Tapi- sis and barringtonensis). nurus to be synonyms ofTropidurus. In Trop- On the mainland, Steindachner (1891) idurus he recognized 11 species: (1) grayii named T. stolzmanni from Peru. Ten years (including bivittata); (2) pacificus; (3) occipi- later, Steindachner (1901) named T. theresiae talis; (4) bocourtii (a new species, synony- from Peru, and subsequently (1902) ampli- mous with occipitalis); (5) peruvianus (in- fied the description. Boulenger (1900) named cluding araucanus, microlophus, heterolepis, Tropidurus thomasi (a junior synonym of T. lessonii, xanthostigma, thoracica, quadrivit- thoracicus) from Peru, and, subsequently in tata, and inguinalis); (6) spinulosus (including 1902 he named T. melanopleurus from Bo- variegatus); (7) torquatus (including tuber- livia. Roux (1907) named T. tschudii (a syn- culata, microlepidotus); (8) hygomi; (9) his- onym of T. occipitalis) from Peru. Muller pidus (including smithii, nigrocollaris, cyclu- (1924) named T. continentalis (a synonym of rus, toelsneri, superciliaris, macrolepis); (10) T. occipitalis) from Ecuador and T. pictus and semitaeniatus; and (11) blainvillii (= Chal- T. praeornatus from Bolivia (both synony- arodon madagascariensis according to Eth- mous with T. melanopleurus). eridge, 1 969a). With the exception ofthe con- Burt and Burt (1930) placed hispida in the troversial status ofPlatynotus (= Tapinurus), synonymy of torquatus, and placed T. tschu- competition between generic arrangements is dii Roux,7T. continentalis Muller, and T. bo- absent after Boulenger's catalog (1885). courti Boulenger into the synonymy of T. oc- Cope (1889) named Tropidurus lemnisca- cipitalis. In 1931 they placed T. hygomi in tus (a junior synonym of T. bivittatus) from the synonymy of T. torquatus. the Galapagos. Subsequently, Baur (1890) Amaral (1933) recognized a new genus, described five new Tropidurus species from Tapinurus, for a new species, scutipunctatus, the Galapagos: albemarlensis, indefatigabilis from Brazil. Schmidt and Inger (1951) con- (= albemarlensis), delanonis, duncanensis, sidered this name to be a junior synonym of and abingdonensis (= pacificus). Boulenger Platynotus semitaeniatus. Recently, Ro- (1891) responded to Cope's (1889) and Baur's drigues (1984b) showed that Platynotus was (1890) treatments of Galapagos Tropidurus a preoccupied name and that Tapinurus by recognizing only three Galapagos species: Amaral (1933) had priority. In addition, Ro- bivittatus (including abingdonensis as a syn- drigues named a new Brazilian species, Tapi- onym), pacificus (including lemniscatus in nurus pinima. Subsequently, Manzini and synonymy), and grayii (including as syn- Abe (1990) described another Tapinurus, T. onyms Baur's albemarlensis, indefatigabilis, helenae. delanonis, and duncanensis). Consistent with Mertens (1956) produced a study of the his view of a multiplicity of species in the Peruvian species of Tropidurus west of the Galapagos, Baur (1892) elevated T. pacificus Andes, recognizing five species in Peru: hol- habelii to species status, supplied an unneed- otropis (= Plica umbra, fide Etheridge, 1970a), ed replacement name for T. delanonis (T. occipitalis, theresiae, peruvianus, and thoraci- hoodensis), and named two new taxa: T. ja- cus. Mertens considered tigris to be a sub- cobii (= T. a. albemarlensis) and T. barring- species of peruvianus, and stolzmanni to be tonensis (= T. albemarlensis barringtonen- a subspecies of occipitalis. He also named T. sis). occipitalis koepekeorum (now T. koepckeo- Heller (1903) added T. grayii magnus (= T. rum) and T. peruvianus salinicola. Donoso- a. albemarlensis) to the list ofdescribed taxa Barros (1966), working in Chile, named five from the Galapagos, and regarded T. bar- new taxa: T. peruvianus maminfensis, T. p. ringtonensis to be a race of grayii and T. marianus, T. p. atacamensis, T. theresioides, abingdonensis Baur to be a synonym of T. and T. tarapacensis. Also, he reduced het- pacificus. erolepis and quadrivittatus to the status of Van Denburgh and Slevin's (1913) revi- subspecies ofperuvianus. Dixon and Wright sion set the current taxonomy of Galapagos (1975) reviewed the Tropidurus of Peru and 8 AMERICAN MUSEUM NOVITATES NO. 3033 revised the arrangement of Mertens (1956). T. divaricatus Rodrigues (1986), which Ro- They recognized that T. occipitalis koepek- drigues (1986) considered to be closely re- eorum and T. o. stolzmanni were distinct spe- lated. The evolution of this group has been cies and also named two subspecies of T. discussed by Rodrigues (1986) and Kasahara thoracicus: talarae and icae. et al. (1987). Ortiz-Zapata (1980b) reviewed the taxonomic status of Tropidurus species in Chile. He synonymized T. peruvianus mamin-ensis MATERIALS AND METHODS with T. theresioides, named T. yanezi, and revalidated the species status of T. quadri- The methodology for formulating general vittatus and T. heterolepis. Subsequently, Or- phylogenetic hypotheses in this study is par- tiz-Zapata (1980c) elevated T. peruvianus simony analysis (Kluge and Farris, 1969; atacamensis to species status and regarded T. Farris and Kluge, 1985, 1986; Farris, 1983; p. marianus as a synonym. Churchill et al., 1985), with its underlying East of the Andes, Roze (1958) named assumption that the weight of all available Tropidurus bogerti from Venezuela. Donoso- evidence must be followed (Kluge, 1989b). Barros (1968) treated it, without discussion, For polarizing hypotheses oftransformation, as a species of Plica, but this view was re- outgroup comparison has been shown to be jected by Etheridge (1970a). the most general method (e.g., Stevens, 1980; Vanzolini and Gomes (1979) started the Farris, 1982; Kluge, 1984, 1985; Brooks and dismemberment of T. torquatus (sensu Burt Wiley, 1985). In particular, a first outgroup and Burt, 1931), by revalidating T. hygomi, ofthe group in question, and the outgroup of of eastern Brazil (although T. hygomi had that combined group are required for ade- earlier been recognized by Etheridge, 1970b, quate transformation polarity (Watrous and without explanation). Cei (1982) followed by Wheeler, 1981; Maddison et al., 1984). For- describing T. etheridgei from Argentina, and tunately, previous studies (Etheridge and de Gudynas and Skuk (1983) named T. cata- Queiroz, 1988; Frost and Etheridge, 1989; lanensis from Uruguay. Rodrigues's (1987) Pregill, in press) have shown that for the study of the species of Tropidurus similar to Tropidurus group the "Stenocercus" group T. torquatus from south ofthe Rio Amazonas ("Stenocercus," "Ophryoessoides," and Proc- went far to sorting the species masquerading totretus) is the first taxonomic outgroup, Lei- under the name T. torquatus. He recognized ocephalus is the second taxonomic outgroup, hygomi, torquatus (including catalanensis as and the Liolaemus group (Liolaemus, Phy- a synonym), etheridgei, hispidus, and de- maturus, and Ctenoblepharys) is the third scribed a number of new species: T. cocoro- taxonomic outgroup (fig. 2). bensis, T. erythrocephalus, T. insulanus. T. The hypothesized character transforma- itambere, T. montanus, T. mucujensis, and tions in this analysis fall into four categories: T. oreadicus. Later, Rodrigues et al. (1988) (1) polarized (additive); (2) unpolarized (ad- recognized another new species, T. psam- ditive); (3) unordered (nonadditive), ancestor monastes, similar to T. hygomi and T. co- hypothesized; (4) unordered (nonadditive), corobensis. ancestor not hypothesized. Polarized trans- Laurent (1982) demonstrated that T. pictus formations may be oftwo characters (ofwhich Muller and T. praeornatus Muller are junior one is hypothesized as ancestral to the other) synonyms of T. melanopleurus, although this or more, but in all cases the additivity of synonymy had been made, without discus- transformatiop is maintained and the ances- sion, by Etheridge (1970b). Rodrigues (1988) tral condition is hypothesized. Unless noted disputed the synonymy of T. pictus (including otherwise, analyzed transformations were T. praeornatus) with T. melanopleurus. polarized (additive). Unpolarized transfor- Surprisingly, three species from Brazil have mations are those in which the ancestral con- come to light that are apparently not closely dition cannot be deduced, but for reason of related to the Tropidurus torquatus group. being either a two-character transformation These are Tropidurus nanuzae Rodrigues or a multicharacter transformation in which (198 1), T. amathites Rodrigues (1 984a), and the order of transformation is hypothesized 1992 FROST: TROPIDURUS GROUP OF LIZARDS 9 on ontogenetic grounds or by reason of mor- phological intermediacy, additivity is main- I 0 tained (i.e., regardless of the ancestral condition it takes two steps to go from character 0 through character 1 to character 2). Unor- dered transformations may have the ancestral condition hypothesized, but additivity is not assumed (i.e., any change from one ho- molog to another is counted as one step), for reason of lack of evidence of any particular polarity between a number ofcharacters. Un- polarized and unordered transformations must be included in the character analysis because the objective of any analysis is to explain all of the data at hand. Additionally, Tropiduridae both ofthese "nontraditional" kinds oftrans- formations have substantial roles to play in Fig. 2. Cladogram of the Tropiduridae. the development of the most parsimonious unrooted network ofterminal taxa. The only difference between these kinds of transformations and standard polarized transfor- native rooting points and character support mations is that they do not necessarily con- for stems. Once sources of cladogram insta- tribute to the polarity vector that determines bility were identified it was possible to dis- rooting of the network to make a tree. cover cladograms unidentified by either an- Occasionally, a particular species could not alytical program. Tree optimization was be assessed for a particular character because obtained using the consistency index (ci) of ofdamage to a specimen or because oflogical Kluge and Farris (1969). Further discussion incongruencies (e.g., shape of a scale in a liz- of analytical techniques can be found under ard lacking that particular scale). In these Results. Only stems supported under some cases, the character was coded as "unknown" character optimization method were consid- for that taxon. Analytical programs used (dis- ered to be supported; no topologies depen- cussed below) allow this by hypothesizing the dent on "empty" stems are discussed. "unknown" to be equivalent to the least re- Because some members of the Tropidurus jected assignment in any particular network group are rare (e.g., Strobilurus torquatus) or topology. in collections not available to me (e.g., T. The transformation series have been ar- psammonastes, T. amathites), I was unable ranged into a data matrix that was subjected to see adequate series of, or, in some cases, to analysis using the PAUP 3.0 (Phylogenetic any specimens ofsome species. Additionally, Analysis Using Parsimony) program ofSwof- many identifications applied to previously ford (1989) and HENNIG86 1.1 (Farris, prepared skeletons (e.g., Tropidurus peruvi- 1988). Because the data matrix is too large anus and T. torquatus) are dubious, given the for evaluating all possible trees or using the recent taxonomic disarray of this group-es- branch-and-bound method of Hendy and pecially if explicit locality data are not sup- Penny (1982), only heuristic methods were plied. Thus, with the exception of a few spe- used to analyze the data. Within PAUP a cies (e.g., T. occipitalis), the amount of number of alternatives of taxon addition, material available was limited. Because of character optimization, and swapping meth- this I have tried to be conservative in char- ods were used. In HENNIG86, extended acterizations. branch-breaking was used. For trees pro- Skeletons, alcoholics, hemipenes, and duced by PAUP, alternative rooting points cleared and double-stained specimens of al- were checked using MacClade (Maddison and most all taxa (see Appendix 1-Specimens Maddison, 1987). The Dos Equis (XX) utility Examined) were examined for interspecific in HENNIG86 was also used to check alter- variation that could be hypothesized to be 10 AMERICAN MUSEUM NOVITATES NO. 3033 apomorphies relative to outgroups (see importance to the systematics of the Tropi- Methods). durus group, but may have some bearing on Members ofthe Tropidurus group that were larger questions of iguanian relationships. In not examined are: polarized transformations below, "0" de- 1. Tropidurus tarapacensis. This is a notes the hypothesized plesiomorphic con- member of the anagenetically tightly knit T. dition and "1" (and higher integers) refers to peruvianus group of species (Ortiz-Zapata, hypothesized apomorphies. In unpolarized 1980a, 1980c). This species is phenotypically and unordered transformations the integer so similar to T. tigris, T. yanezi, and T. ther- assignment is arbitrary. esioides that its absence probably has little Throughout this discussion I will use the effect on the analysis. following collective terms (species not ex- 2. Tropidurus psammonastes. This is one amined noted with a t): of the recently described species (Rodrigues (1) Tropidurus occipitalis group: Tropi- et al., 1988) ofthe Tropidurus torquatus group durus grayii complex (T. albemarlensis, T. from Brazil that is, according to the original duncanensis, T. delanonis, T. grayii, and T. description, closely similar to T. hygomi. Ad- pacificus), T. bivittatus, T. habelii, T. koepek- ditionally, although I had access to alcoholic eorum, T. occipitalis, and T. stolzmanni. material of T. cocorobensis, T. insulanus, and (2) Tropidurus peruvianus group: Tropi- T. mucujensis (Rodrigues, 1987) 1 was unable durus atacamensis, T. heterolepis, T. peru- to prepare skeletons or hemipenes of these vianus, T. quadrivittatus, tT. tarapacensis, T. (although X-rays were available). theresiae, T. theresioides, T. thoracicus, T. 3. Tropidurus amathites Rodrigues (1984b) tigris, T. yanezi. and T. divaricatus Rodrigues (1986). These (3) Western Tropidurus group: Tropidurus two recently described Brazilian species pre- occipitalis group + T. peruvianus group. sumably are close to T. nanuzae Rodrigues, (4) Eastern Tropidurus group (or Tropi- 1981, sharing with it a reduced sternal fon- durus group east of the Andes)4: tTropidurus tanelle (Rodrigues, 1986) and a presumably amathites, tT. divaricatus, T. torquatus group derived karyotype (Kasahara et al., 1987). (see below for content), T. melanopleurus, T. Like the recently described Brazilian species nanuzae, T. spinulosus, Plica, Strobilurus, in the Tropidurus torquatus group, T. psam- Tapinurus, Uracentron, Uranoscodon. monastes, these were not available to me. (5) Tropidurus torquatus group: T. bogerti, 4. Tapinurus helenae Manzini and Abe T. cocorobensis, T. erythrocephalus, T. eth- (1990). This Brazilian species is clearly close- eridgei, T. hispidus, T. hygomi, T. insulanus, ly related to the other highly apomorphic spe- T. itambere, T. montanus, T. mucujensis, T. cies of Tapinurus, and appears to differ from oreadicus, tT. psammonastes, T. torquatus. Tapinurus semitaeniatus only in color pattern and minor aspects of squamation. CRANIAL CHARACTERS 1. Skull size: (0) adult males with head TRANSFORMATION SERIES length < 23 percent of snout-vent length; (1) adult males with head length > 23 percent In this section I will neither give an ex- of snout-vent length. Members of the Tropi- haustive summary of the morphology and durus group east ofthe Andes, excluding Ur- anatomy ofmembers ofthe Tropidurus group, anoscodon, have relatively large heads, par- nor will I solely summarize variation that can ticularly when compared to the Tropidurus be characterized successfully in transforma- group west of the Andes. The "Stenocercus" tion series. Rather, I will present a brief sum- group and species of Leiocephalus also have mary ofthe features that can be characterized relatively small heads. in ways that allow phylogenetic inferences to be drawn; those that cannot be characterized 4I recognize that the eastemmost "western" Tropi- successfully now, but that may be of interest durus, T. stolzmanni, is found east of the continental to successive systematists, are noted where divide in the Huancabamba Depression Region. How- appropriate. Additionally, I will occasionally ever, this generalization of "eastern" and "western" is digress into areas that may be of no great so convenient that I overlook the geographical anomaly. 19921992FROST: TROPIDURUS GROUP OF LIZARDS I1I

2. Skull elevation (fig. 3): (0) skull not elevated at level of orbits (skull height < 39% ofskull length) -postorbital bone not rotated to form a flange; (1) skull elevated at level of orbits (skull height > 39% of skull length)- postorbital bone rotated outward to form a flange. Uranoscodon superciliosus and Plica umbra have noticeably elevated skulls compared with the remainder of the Tropidurus group and the outgroups. This elevation is correlated with the enlargement ofthe orbits and concomitant rotation of the postorbital bones to form postorbital flanges. Although some members of the outgroups (e.g., some "Ophryoessoides") show moderate elevation, these species are removed phylogenetically from the base of the "Stenocercus" group; therefore, the similar condition is considered homoplastic. 3. Skull compression (fig. 3): (0) not compressed- skull height > 30 percent of skull length; (1) compressed-skull height < 25 percent of skull length. Tapinurus shows extreme dorsoventral compression of the skull (and body) relative to all other members of Fig. 3. Lateral views ofskulls. A: Plica umbra, the Tropidurus group and outgroups. KU 146659, showing great skull elevation and 4. Rostrum length (fig. 3): (0) long; (1) shortening ofrostrum. B: Tropidurus atacamensis, shortened. Within the Tropidurus group, only KU 161986, showing no pronounced elevation of Plica umbra and Uranoscodon superciliosus skull or shortening ofrostrum. C: Tapinurus semitaeniatus, LSUMZ 39519, showing great skull show any perceptible shortening of the ros- compression. Scales = 5 mm. Arrows show en- trum (preorbital length of the skull). In both larged maxillary nutritive foramina. of these species this may be correlated with elevation ofthe skull in the orbital region. A related characteristic, rostrum shape, has de- fied adequate characterization. Upon casual 5. Premaxilla (fig. 4): (0) nasal spine nar- inspection, Uranoscodon appears to have a row-dentigerous part of premaxilla broad; blunt rostrum. What causes this illusion is (1) nasal spine broad-dentigerous part of the extreme elevation ofthe skull at the level premaxilla narrow. In Plica, Strobilurus, ofthe orbits; Uranoscodon has an acute snout. Uracentron, and Uranoscodon the dentiger- Only individuals of Plica exhibit a blunt ous part of the premaxilla is narrowed with snout. However, Plica lumaria does not ex- respect to the remaining Tropidurus group hibit this feature and P. plica is sexually di- and outgroups. One would expect this feature morphic in this character. Males (e.g., MAN to be highly correlated with the number of 76, MCZ 43865) have a snout relatively more premaxillary teeth a priori, but this is not so; acute than females. Only female Plica plica see Transformation Series 19. Although aut- (e.g., AMNH 61314, KU 167499) could be apomorphic, T. spinulosus has a particularly coded as having blunt snouts. However, the narrow nasal spine relative to all other mem- fact that this is sexually dimorphic in Plica bers of Tropidurus east of the Andes. plica, absent in the almost identical P. lu- 6. Nasal bones (fig. 5): (0) not reduced- maria, and not sexually dimorphic in Plica external choana does not approach level of umbra makes me suspicious that these fea- anterior extent of prefrontal; (1) reduced- tures are not homologous, and, therefore, I excavated to a point approaching the anterior have not used them in my analysis. part of the prefrontal in T. bogerti; (2) ex- 12 AMERICAN MUSEUM NOVITATES NO. 3033

A .. . . . :

Fig. 4. Anterior views ofpremaxilla. A: Tropi- Fig. 5. Dorsal views of skulls. Nasal bones are durus melanopleurus, KU 136370. B: Plica plica, shaded. A: Tapinurus semitaeniatus, LSUMZ MAN 76. Scales = 1 mm. 39519, extreme excavation of nasals. B: Tropi- durus bogerti, RWM 11662, some excavation of nasals. C: Tropidurus stolzmanni, KU 134747, no excavation of nasals. Scales = 10 mm. cavated to a point where the prefrontal contacts the margin of the external choana in Tapinurus. Although short-snouted mem- 7. Nutritive foramina ofmaxillary (fig. 3): bers ofthe Tropidurus group (i.e., Plica, Ura- (0) small, inconspicuous; (1) enlarged, con- noscodon) appear upon cursory glance to have spicuous. The characteristic of greatly en- the nasal bones reduced, more careful ex- larged nutritive foramina along the lateral amination shows that this is not the case. surface of the maxilla is unique among igua- This is an illusion caused by the extreme hy- nians and characteristic of the Tropidurus pertrophy in size ofthe orbits in these species. group east of the Andes, excluding Uranos- Only Tropidurus bogerti and Tapinurus (most codon. extreme) show any retreat of the nasals. 8. Maxillopalatine foramen (infraorbital 1992 FROST: TROPIDURUS GROUP OF LIZARDS 13

foramen

lacrimal

maxillopalatine foramen Fig. 6. Posterior view ofanterior margin ofleft orbit showing maxillopalatine foramina. Left: Tropidu- rus atacamensis, KU 161986. Right: Plica plica, MAN 76. Scales = 1 mm. canal ofJollie, 1960) (fig. 6): (0) much smaller mosal bone curved around the posterior end than lacrimal foramen; (1) enlarged and dor- of the temporal fenestra-the posterior apex soventrally expanded-frequently subequal of the temporal fenestra forming a smooth in size to lacrimal foramen. In the outgroups curve. This is reflected in shape changes of (as well as in most iguanians) the maxillo- the temporal fenestra in the three species of palatine foramen is considerably smaller than Plica. the lacrimal canal. However, in Plica, Ura- 11. Superior fossa of quadrate (fig. 8): (0) centron, and Strobilurus the maxillopalatine relatively small, a process of the squamosal foramen is usually considerably enlarged dor- fitting into the fossa like a peg in a hole; (1) soventrally and as large as, or larger than, the relatively enlarged, the squamosal not pen- lacrimal foramen. Some individuals of T. etrating the fossa. In the Tropidurus group koepckeorum, T. hispidus, and T. hygomi the superior fossa ofthe quadrate is enlarged. show somewhat enlarged maxillopalatine fo- The functional significance is unknown; how- ramina that are expanded laterally but I do ever, as noted elsewhere, members of the not regard this as a homologous condition. Tropidurus group are distinctive within the 9. Pineal foramen: (0) present; (1) absent. iguanians for their general delicateness ofbone Although some members of the "Stenocer- structure. The increase in size of the fossa cus" group lack the pineal foramen (e.g., and the failure of the squamosal to develop "Stenocercus" empetrus), these species are enough of a quadrate process to fit into the removed from the "base" of "Stenocercus," hole may well be a result of this trend. The so this similarity is likely homoplastic with "Stenocercus" group and Leiocephalus show respect to this feature in the Tropidurus group. some approach to this condition relative to Within the Tropidurus group this is an aut- the remaining iguanians but even in these a apomorphy of Uracentron azureum (but process of the squamosal penetrates into the present rudimentarily in one specimen of U. superior fossa of the quadrate. azureum examined, AMNH 60330). 10. Squamosal shape and skull width (fig. 7): (0) squamosal bone relatively straight, re- MANDIBULAR CHARACrERS flected in the posterior apex of the temporal 12. Alveolar shelf of mandible (fig. 9): (0) fenestra forming an acute angle; (1) squa- forming a well-defined ridge; (1) alveolar ridge 14 AMERICAN MUSEUM NOVITATES NO. 3033

A quadrate

- anterior

Fig. 8. Squamosal-quadrate articulations. Fig. 7. Dorsal views of skulls. A: Tropidurus A: Leiocephalus carinatus, UMMZ 149104; su- stolzmanni, KU 134747. B: Plica umbra, KU perior fossa ofquadrate small. B: Tropidurus bog- 146659. Scale = 10 mm. erti, RWM 11662; superior fossa of quadrate enlarged. Scales = 2 mm. rounded-erosion of thickness of mandible below level ofalveolar ridge; (2) alveolar ridge there is very weak overlap and members of poorly defined- medially approaches ventral the Tropidurus group east of the Andes (ex- margin of mandible. In outgroups, the alve- cept Uranoscodon) tend to have stronger de- olar shelf of the mandible is well defined. velopment of the process than members of However, in the Tropidurus group, the al- the Tropidurus group west of the Andes. veolar shelf is slightly to greatly eroded. In However, this is to considerable Uranoscodon subject vari- and Tropidurus west ofthe An- ation, which prevents me from hypothesizing des, the degree of erosion is not as great as a shift from weak development to strong de- that seen in the remaining Tropidurus group. velopment as a transformation series. 13. Lingual coronoid process of dentary 14. Posterior extent ofdentary (fig. 10): (0) (fig. 9): (0) not overlapping anterior lingual dentary extending < 50 percent ofthe length "leg" of coronoid; (1) overlapping anterior from apex of coronoid to anterior edge of lingual "leg" of coronoid. Unique among articular; (1) extending > 50 percent of the iguanians examined, members of the Tropi- length from the apex of the coronoid to the durus group, excepting Uranoscodon, exhibit anterior edge of the articular. Excluding Ur- a process ofthe dentary that overlaps part of anoscodon, the the anterior Tropidurus group east of the labial "leg" of the coronoid. In Andes exhibits a more posterior extension of some members of the "Stenocercus" group the dentary than the Tropidurus group west 1992 FROST: TROPIDURUS GROUP OF LIZARDS 15

B il

2 Fig. 10. Labial views of dentaries. A: Uranos- codon superciliosus, REE 16600. B: Tropidurus bogerti, RWM 11662. C: Tropidurus melanopleu- Fig. 9. Lingual views ofmandibles. A: "Steno- rus, KU 136367. Scales = 5 mm. Shown: (1) ex- cercus" apurimacus, KU 134284. B: Tropidurus tensive posterior extent of dentary; (2) capture of stolzmanni, KU 134747. C: Plica plica, MAN 76. anterior surangular foramen between coronoid and Shown: (1) weakly eroded alveolar shelf; (2) strongly dentary. eroded alveolar shelf; (3) lingual coronoid process of dentary. Scales = 2 mm. 16. Angular condition (fig. 1 1): (0) large, distinct; (1) reduced (limited to below sur- ofthe Andes or the outgroups ("Stenocercus" angular-prearticular suture) or lost. Uranos- group and Leiocephalus). Some Leiocephalus codon and Tropidurus from west ofthe Andes species approach "1" but they still lack the have relatively large angulars identical to condition as seen in the Tropidurus group. those in the outgroups. East ofthe Andes, all 15. Anterior surangular foramen (fig. 10): members ofthe Tropidurus group (excluding (0) not captured by contact of the coronoid Uranoscodon) show some reduction in size and dentary posterior to the foramen; (1) of the angular, but inter- and intraspecific "captured" by contact of the coronoid and variation in shape and presence or absence surangular posterior to the anterior suran- is confusing; it will be discussed here but not gular foramen. Excepting Uranoscodon and placed in any transformation series. Ofthose Tropidurus bogerti, the Tropidurus group east with a reduced angular, some Tapinurus of the Andes is characterized by enclosure of semitaeniatus (MCZ 79805) and T. spinulo- the anterior surangular foramen between the sus show the least reduction of the angular; coronoid and dentary. Leiocephalus also has it is only moderately reduced from the plesio- the anterior surangular foramen enclosed by morphic condition. However, in some other posterior contact of the coronoid and den- T. spinulosus (e.g., CAS 49843) the angular tary, but the "Stenocercus" group and the apparently is fused with the prearticular and Tropidurus group west ofthe Andes lack this not visible. In Plica plica and P. lumaria the contact. Outgroup comparison is insufficient angular and splenial apparently are fused and to polarize this transformation series; there- this composite bone contacts the surangular, fore it is treated as unpolarized (i.e., the "an- although in MCZ 85313 (Plica plica) there is cestor" is coded as "unknown") in this anal- a slight separation ofthe splenial and angular. ysis. In Plica umbra, most Tapinurus semitaenia- 16 AMERICAN MUSEUM NOVITATES NO. 3033

prearticular (T. hispidus, T. itambere) or closely approach it (T. erythrocephalus, T. etheridgei, T. hygomi, T. melanopleurus, T. nanuzae, T. torquatus, and Strobilurus torquatus) although characterization of this transformation would be difficult because of intraspecific variation. 17. Posterior mylohyoid foramen, osseous contact (fig. 11): (0) posterior mylohyoid foramen penetrating angular or between dentary and angular; (1) between angular and splenial; (2) between dentary and splenial. In outgroups and the Tropidurus group west of the Andes (and Uranoscodon) the anterior mylohyoid foramen is invariably in contact with the angular. Within this category several conditions obtain that cannot be polarized because of variation in the outgroups. Most of the T. occipitalis group and the T. peruvianus group show the posterior mylohyoid foramen either penetrating the angular or penetrating between the dentary and angular. However, T. habelii and T. bivittatus have the posterior mylohyoid foramen penetrating between the angular and splenial, presumably an apomorphy uniting these species. All of the Tropidurus group east of the Andes, except for Uranoscodon, T. spinulosus, and Uracentron, exhibit a condition where the posterior mylohyoid foramen is excluded from contact with the angular (or even the topographic "angular region" when the angular is absent). Fig. 11. Ventral views of right mandibles: an- 18. Posterior mylohyoid foramen, posi- gulars and posterior mylohyoid foramina. A: Ura- tion (fig. 11): (0) at the level of anterior end noscodon superciliosus, REE 16600. B: Tropidurus of mandibular fossa; (1) placed more poste- bivittatus, LACM 10637. C: Tropidurus spinulo- riorly, about 33 percent of the length of the sus, USNM 125166. D: Tropidurus melanopleu- mandibular fossa back from the anterior end. rus, KU 136367. E: Uracentron flaviceps, KU The position ofthe mylohyoid foramen var- 175317. F: Plica plica, MAN 76. Scales = 2 mm. ies considerably in iguanians and may be of Nominal angulars are stippled. Nominal splenials are vertically screened. Posterior mylohyoid fo- wide systematic application. In the outgroups ramina denoted with arrows. (Leiocephalus and "Stenocercus" group) and most members of the Tropidurus group, the posterior mylohyoid foramen is on the ventral surface of the dentary at the level of the tus, Tropidurus bogerti, and Uracentron, the anterior end of the adductor fossa. In Ura- angular is present as a small flake of bone centron and Tapinurus, however, it is dis- that actually may be a dissociated piece of placed considerably posteriorly. In MCZ the splenial. In the remaining Tropidurus 172948 (T. erythrocephalus) the posterior group the angular is absent or is fused either mylohyoid foramen is displaced posteriorly, to the splenial or prearticular. In this group though not to the extent found in Uracentron the splenial can be widely separated from the and Tapinurus. 1992 FROST: TROPIDURUS GROUP OF LIZARDS 17

DENTITION 19. Premaxillary teeth, number (fig. 4): (0) 6-7; (1) 4-5. In most ofthe Tropidurus group, as well as the outgroups, the premaxilla normally bears 6-7 teeth. In Uracentron, Strobi- lurus, Tapinurus, Uranoscodon, Plica plica, and P. lumaria, this number is reduced to 5 A or 4. The trend is for those species with narrow premaxillae (see Transformation Series 5) to have fewer premaxillary teeth, but this correlation is not perfect. Although Plica umbra is coded as "0", the condition may not be homologous with other "0" conditions, inasmuch as the medial premaxillary teeth in Plica umbra are frequently very small, allow- ing the conjecture that they are in a sense "supernumerary." 20. Anterior maxillary teeth, enlargement (Boulenger, 1885; Etheridge, 1970a) (fig. 3): (0) not or only feebly enlarged in older adults; (1) enlarged in older adults, forming canini- form teeth. Within the Tropidurus group many species show ontogenetic enlargement ofthe anteriormost few maxillary teeth, concomitant with an upward tilting ofthe lower margin ofthe premaxilla, resulting in the ap- some evi- Fig. 12. Posterior maxillary dentition. A: Lei- pearance of "canines." There is ocephalus carinatus, UMMZ 149104. B: Uranos- dence of this trend in Leiocephalus (G. Pre- codon superciliosus, REE 16600. C: Tropidurus gill, personal commun.) and some species in melanopleurus, KU 136367. D: Plica umbra, KU the "Stenocercus" group (e.g., "S." festae, 146659. E: Plica plica, MAN 76. Scale = 1 mm. Proctotretus pectinatus) show ontogenetic elevation of the premaxilla without concomitant lengthening of anterior maxillary teeth. Premaxillary elevation such as this is wide- 21. Posterior maxillary and dentary teeth, spread outside of the Tropidurinae, such as crown flaring (Etheridge, 1966, 1968, 1970a; in some Crotaphytus (Crotaphytidae), Cteno- Etheridge and de Queiroz, 1988) (fig. 12): saura (Iguanidae), and Phymaturus (Tropi- (0) shaft parallel-sided with crowns not or duridae: Liolaeminae), and it cannot be char- weakly flared; (1) crowns flared. The shape acterized or polarized in any satisfactory way. of the crowns of cheek teeth is difficult to However, all species ofthe Tropidurus group describe and is beset with outgroup compar- east of the Andes (except Uranoscodon and ison problems. Leiocephalus has flared teeth, some individuals of Tropidurus melanopleu- but the "Stenocercus" group exhibits all con- rus [taxon coded as "1" because present in ditions from weakly flared and tricuspid (e.g., older individuals]) exhibit "canine" forma- "S." guentheri) to peglike with poorly de- tion well advanced over that seen in the out- veloped cusps (e.g., Proctotretus pectinatus). groups or the species of Tropidurus found The variation among conditions within the west ofthe Andes. Greatest development may Tropidurus group is necessarily nonpolar- be in Plica umbra, although this is ap- ized. proached in Uracentronflaviceps. Small sam- Beyond the nonflared-flared dichotomy, ple size precludes the recognition ofmultiple characterization ofthe tooth shape of species characters within this transformation series. in the Tropidurus group is difficult to assess 18 AMERICAN MUSEUM NOVITATES NO. 3033

oftooth flaring as a diagnostic feature uniting Plica, Strobilurus, and Uracentron, and, as a modality (i.e., in Plica umbra and Uracen- tron azureum), I agree. Tapinurus and Tro- pidurus hispidus also have (predominantly) flared teeth. 22. Posterior maxillary teeth, elevations: (0) posterior maxillary teeth apparently hyp- sodont, extending above the edge ofthe maxilla more than the width of a tooth; (1) posterior maxillary teeth brachydont, not extending far above the level of the maxilla. In Tropidurus nanuzae, Uracentronflaviceps, and Plica a crest of bone along the ventral edge ofthe maxilla deepens the dental gutter, making the teeth appear shorter when viewed laterally. 23. Posterior maxillary teeth, orientation (fig. 13): (0) posterior maxillary teeth set obliquely on the maxilla; when viewed from the ventral side most of the length of the individual teeth can be seen; (1) posterior maxillary teeth set more vertically on maxilla (not to be confused with recurving ofthe teeth as seen in some other taxa such as some Lei- ocephalus); when viewed from the ventral side, much ofthe length ofthe tooth is hidden from view; additionally, the orbital margins Fig. 13. Ventral views of skulls. A: Tropidurus ofthe jugal form "cheeks" that, as evidenced stolzmanni, KU 134747. B: Plica umbra, KU by their outward rotation, are structurally part 146659. Scales = 10 mm. of the inward rotation of the dental row. In some members ofthe Tropidurus group (i.e., Uranoscodon, Plica, Tropidurus nanuzae, and essentially continuous. I will discuss the Strobilurus, and Uracentron) the jugal and species as if they fall into four groups; the maxilla are exceptionally broad in the region distinctions among them are fuzzy. Within of the posteriormost teeth (reflected also in the Tropidurus group, only Uranoscodon (fig. Transformation Series 22). Uranoscodon, 12) and Tropidurus theresiae show constric- Plicaplica, and P. lumaria differ slightly from tion of the crown of the tooth relative to the the other taxa coded as apomorphic in this shaft. A larger group exhibits posterior regard by not having the teeth set quite so maxillary teeth that are, or are nearly, par- vertically. None of the remaining species in allel-sided: T. bogerti, T. cocorobensis, T. er- the Tropidurus group, nor outgroups, exhibits ythrocephalus, T. insulanus, T. itambere, T. this condition. melanopleurus (fig. 12), T. montanus, T. 24. Pterygoid teeth (Etheridge, 1966): (0) nanuzae, T. spinulosus, T. torquatus, and all present; (1) absent. Although several species of the T. occipitalis group and T. peruvianus of Leiocephalus lack pterygoid teeth, other group, except T. theresiae. Of these, T. bivit- species phylogenetically more "basal" in that tatus approaches the condition of being genus have them (e.g., L. carinatus) (Pregill, slightly flared. Some Tapinurus semitaenia- in press); therefore I have considered Lei- tus, some Plica umbra (fig. 12), some Ura- ocephalus to plesiomorphically have ptery- centron azureum, Tropidurus hygomi, and T. goid teeth. "Stenocercus" group members spinulosus could be called slightly flared. Eth- have pterygoid teeth, with the exception in eridge (1966, 1968, 1970a) used the presence my material of some "S." nigromaculatus 1992 FROST: TROPIDURUS GROUP OF LIZARDS 19 -c D

Fig. 14. Pectoral girdles and ribs. A: Leiocephalus schreibersi, KU 93358. B: Tropidurus stolzmanni, KU 134704. C: Tropidurus hispidus, KU 167513. D: Tropidurus bogerti, RWM 11663. E: Tapinurus semitaeniatus, LSUMZ 39519. Scales = 5 mm. and several members ofthe derived "S." hu- vicular fenestra if they had a place to put it. meralis group (e.g., "S." boettgeri, "S." cras- Therefore, clavicular fenestration cannot be sicaudatus, and "S." humeralis). Presence is placed with any assuredness at any particular therefore considered the plesiomorphic con- level of universality. dition within the Tropidurus group, in which 26. Sternum, fenestration (Rodrigues, Tropidurus bogerti, Plica umbra, Strobilurus 1986) (fig. 14): (0) single fenestration present; torquatus, Tapinurus semitaeniatus, and (1) fenestra absent. With the exception of Uracentron azureum lack pterygoid teeth. Tropidurus nanuzae (and T. divaricatus and T. amathites, according to Rodrigues, 1986 [not seen by me]) all members of the Tropi- PEcroRAL GIRDLE durinae have medially fenestrate sterna, 25. Clavicle (Etheridge and de Queiroz, which otherwise may be a synapomorphy of 1988) (fig. 14): (0) strongly flanged, frequently Tropiduridae and Phrynosomatidae (Frost fenestrate; (1) weakly flared or cylindrical, and Etheridge, 1989). Sternal fontanelles are never fenestrate. In the outgroups, all of the also absent in some members of the Liolae- Tropidurus group west of the Andes, and, of minae, including Liolaemus occipitalis (Kel- those east ofthe Andes, Tropidurus nanuzae, ler and Krause, 1986) as well as several other Plica umbra, Uranoscodon, and Uracentron, species (R. Etheridge, personal commun.). the clavicle has a well-developed postero- 27. Posterior process of the interclavicle ventral blade that approaches the lateral pro- anterior to the sternum (fig. 14): (0) "free" cesses of the interclavicle. Predominantly in part ofthe posterior process of the interclav- these, the flange is penetrated by a fenestra. icle > 25 percent of the total length of the In some Tropidurus occipitalis (e.g., KU sternum (i.e., the sternum is small); (1) "free" 142714 and 142721) and in Uracentronflavi- part ofthe posterior process ofthe interclav- ceps these fenestrae are not present. Plesio- icle < 25 percent of the total length of the morphically within the "Stenocercus" group, sternum (i.e., the sternum is enlarged). Among the clavicle is flanged, although it is uncom- iguanians, only the Phrynosomatidae and the monly fenestrate in adults (as in "S." praeor- Tropidurus group show this apomorphic natus). Leiocephalus also has a weakly flanged manifestation ofenlargement ofthe sternum. clavicle, so, within the Tropidurus group this Tropidurus spinulosus most closely ap- condition must be considered plesiomorphic. proaches the plesiomorphic condition within Whether the presence of a clavicular fe- the Tropidurus group at 24 percent. Most oth- nestra is an attribute useful for this phylo- er species are from 12 to 19 percent with T. genetic reconstruction is arguable. Species that nanuzae at 0 percent. do not have a clavicular flange cannot be as- Other aspects of the interclavicle require sessed as to whether they would have a cla- some discussion. Across the Iguania varia- 20 AMERICAN MUSEUM NOVITATES NO. 3033

plica, and Uracentron), but a few show extension far beyond the primary coracoid ray (i.e., Tapinurus semitaeniatus, Tropidurus itambere, some Plica plica, T. melanopleurus). Further work on this character may prove fruitful. 28. Interclavicle median process (Ether- idge, 1964; Etheridge and de Queiroz, 1988) A B C (fig. 14): (0) posterior process of the interclavicle extending as a broad process poste- Fig. 15. Lateral view of right scapulocora- riorly well beyond the posterolateral corners coids. A: Tropidurus albemarlensis, LACM of the sternum; (1) posterior process of the 196261. B: Tropidurus spinulosus, KU 97856. C: interclavicle not extending posteriorly be- Tapinurus semitaeniatus, LSUMZ 39519. (1) yond the posterolateral corners of the ster- scapular fenestra; (2) suprascapular fenestrae. num. In the Tropidurus group, except for Tropidurus bogerti, T. spinulosus, and Ura- tion in the angle made by the lateral and pos- centron azureum, the posterior process ofthe terior processes ofthe interclavicle is contin- interclavicle stops short of the level of the uous and extremely difficult to characterize. lateral corners of the sternum. In both the However, within this restricted set ofgroups, "Stenocercus" group and Leiocephalinae, the characterization proved difficult only within posterior process ofthe interclavicle extends the T. peruvianus group; additional speci- well past this level. mens examined may show that I have erred 29. Scapular deflection and fenestration of in my evaluation of these species. Most of scapulocoracoid (Lecuru, 1968; Etheridgeand the "Stenocercus" group have anchor-shaped de Queiroz, 1988) (fig. 15): (0) scapular fe- interclavicles, the angle varying between 70 nestra present, large, and scapula not bent; and 750; Leiocephalus is somewhat more (1) scapular fenestra present, reduced, scap- acute, around 65-70°. The modal interval ula weakly bent; (2) scapular fenestra absent, within the Tropidurus group is similar to that with no room for fenestra in scapula because in the "Stenocercus" group plesiomorphical- extremely bent. In some Tropidurus (e.g., T. ly, but about 800 in the T. peruvianus group bogerti, T. melanopleurus, and T. spinulosus) and, except for T. nanuzae (650), the Tropi- the scapula is deflected noticeably inward, durus group east of the Andes. Tapinurus is causing the scapulocoracoid to form a more exceptional in that the angle formed is greater acute bend than normally seen in outgroup than 1000. Unfortunately, interspecific vari- species. Concomitantly, this bend reduces the ation is extensive enough that it overwhelms size of the scapular fenestra to the point that any attempt to recognize discrete steps for it is absent as an individual variation in T. analysis. melanopleurus and T. spinulosus. Tapinurus A related character, length of lateral pro- shows an even more apomorphic condition; cesses of the interclavicle, also proved too in this case the scapulocoracoid is bent al- variable for use in a transformation series. most in half, with no room for the scapular Most species have the lateral processes not fenestra at all. Other lizards that live in cracks extending to the medial portion of the pri- in rocks have a strongly inflected shoulder mary coracoid ray (i.e., Leiocephalus, most girdle, but in Phymaturus (Liolaeminae) and "Stenocercus" group, T. grayii complex, T. Sauromalus (Iguanidae) the inflection is at bivittatus, most T. peruvianus group, T. the scapula-suprascapula suture (R. Ether- koepekeorum, some T. occipitalis, T. stolz- idge, personal commun.). manni, some T. etheridgei, some Plica um- 30. Suprascapular fenestrations (fig. 15): bra, and Uranoscodon), or extending only to (0) absent, or very tiny; (1) large. Within the the medial extent of primary coracoid ray suprascapula in species of the Tropidurus (i.e., T. habelii, T. atacamensis, some T. oc- group east of the Andes (excluding Uracen- cipitalis, T. bogerti, some T. etheridgei, T. tron), nonossified, nonchondrified fenestrae hispidus, T. hygomi, T. torquatus, some Plica appear. Within the "Stenocercus" group I 1 9921992FROST: TROPIDURUS GROUP OF LIZARDS 21 have seen these fenestrae only in "Stenocer- cus" praeornatus, which is not phylogenetically near the "base" of the "Stenocercus" group (i.e., it is in a derived monophyletic group containing, among others, "S." humeralis and "S." empetrus). I have not seen these fenestrations within Leiocephalus. Tropidurus koepckeorum and T. occipitalis may have tiny fenestrations along the suprascapular margin as an individual variation. Strictly speaking these "fenestrations" are zones of connective tissue that lack either chondrification or ossification and are not actually holes, but appear as clear windows in - - )~~~~~I double-stained specimens. 31. Rib formula (fig. 14): (0) five ribs in contact with the sternum and xiphisternum: 3 sternal ribs + 2 xiphisternal ribs; (1) six ribs in contact with the sternum and xiphisternum (3 sternal ribs + 3 xiphisternal ribs, Fig. 16. Left humeri. A: Uranoscodon super- or 4 sternal ribs + 2 xiphisternal ribs), with ciliosus, MCZ 58340. B: Tropidurus peruvianus, the insertion of the fourth sternal rib being SDSNH 30843. C: Tropidurus melanopleurus, KU very close to the insertion ofthe xiphisternal 136367. All scales = 10 mm. rods or conversely in the case of only three sternal ribs, the first xiphisternal rib inserting very close to the insertion ofthe xiphisternal ciated with entire origin of the pectoralis rods on the sternum. In most species and the musculature. Tapinurus, unique among igua- outgroups, there are three sternal ribs and two nians examined, has long cartilaginous rods xiphisternal ribs. Leiocephalus is apomor- in that follow the origin of the m. pectoralis to phic that the second xiphisternal rib bears form recurved bars that pass from connection a posteriorly directed and recurved rod. In Tropidurus atacamensis, T. bogerti, T. with the xiphisternum ventrally across the T. posteriormost xiphisternal ribs. Short exten- hispidus, itambere, T. montanus, T. tor- sions from the postxiphisternal bars, asso- quatus, T. melanopleurus, T. spinulosus, Pli- ciated with ventral origin ofthe m. pectoralis, ca plica, P. lumaria, Strobilurus torquatus, can also be seen in most specimens of most and Uracentron there are six total sternal + species of the Tropidurus group east of the xiphisternal ribs. The fourth in the series can Andes (excluding Uranoscodon) and, albe attached, as an individual variation, either though not coded as a transformation be- very near to the xiphisternal bar on the ster- cause ofintraspecific variation, is likely a syn- num or very near the sternum on the xiphi- apomorphy of this more inclusive group. sternal bar. In UMMZ 129418 (Uracentron Leiocephalus has superficially similar (non- azureum) the last xiphisternal rib is anom- homologous) bars that are not associated with alously reduced to a free bar attached only to the the pectoralis musculature and pass dorsally xiphisternal bar. Plica umbra and Ura- to the posteriormost xiphisternal ribs. centron azureum share the condition of the sternum and postxiphisternal inscriptional bars being elongated posteriorly, making the LIMBS postxiphisternal elements appear "stretched" 33. Humerus, head (fig. 16): (0) articular posteriorly. surface scroll-like; (1) somewhat elevated, 32. Recurved xiphisternal-pectoral ribs ovate; (2) ball-shaped and very elevated. (fig. 14): (0) absent or present as short spurs Members ofthe Tropidurus group, excluding associated with the medial part of the pec- Uranoscodon, effectively have a ball-and- toralis musculature; (1) present, long, asso- socket shoulderjoint, which is, to my knowl- 22 AMERICAN MUSEUM NOVITATES NO. 3033

finger relative to the third. This elongation is primarily the result ofelongation ofthe fourth metacarpal to approximate equal length with the third metacarpal. In Uranoscodon, Tropi- durus west of the Andes, and the outgroups, the fourth metacarpal is distinctly shorter than the third. A subset ofthe elongate fourth metacarpal group has continued elongation ofthe fourth finger by lengthening the first phalanx. Tropi- durus bogerti, T. spinulosus, T. melanopleurus, Plica, Strobilurus, Tapinurus, and Ura- Fig. 17. Dorsal views of left hands. A: Tropi- centron. Tropidurus etheridgei, T. hygomi, and durus bivittatus, LACM 106307; arrow points to T. torquatus approach this condition. medial centrale; fourth metacarpal (stippled) 36. Claw of first toe: (0) weakly flexed; (1) shorter than third metacarpal; first phalanx of strongly flexed, recurved. Plica, Strobilurus, fourth finger (cross-hatched) distinctly shorter than and some Tropidurus show some degree of first phalanx of third finger. B: Tropidurus hispi- recurving of the claw of the first toe. Only in dus, KU 167508; medial centrale absent; fourth Uracentron, however, can this feature be metacarpal elongated; first phalanx offourth finger characterized adequately. As noted by Eth- not elongated. C: Uracentronflaviceps, KU 175317; eridge (1 970a), another feature, digits bent medial centrale absent; fourth metacarpal elon- to gate; first phalanx elongate. strongly at their articulations (they appear be "physically challenged"), is a characteristic of Plica and some Tropidurus. I found this feature widespread within species east of edge, unique among lizards. Members of the the Andes, but impossible to characterize Tropidurus group west ofthe Andes have the across all taxa, although there is increased ball-and-socket less well developed than in incidence of, and maximal development in, Tropidurus east of the Andes, Strobilurus, some specimens of T. spinulosus, Plica, and Plica, and Uracentron. Leiocephalus is here Uracentron. regarded as having the plesiomorphic con- 37. Fringe on fourth toe: (0) absent; (1) dition, but it is apomorphic in its own way present. Unique within the Tropidurus group by having a lateral rotation ofthe head ofthe and the outgroups, Uranoscodon has the scales humerus. of the edges of the fourth developed into 34. Medial centrale (fig. 17): (0) present; fringes, much as in Basiliscus (Corytophan- (1) absent. The loss of the medial centrale is idae). unique among iguanians. All species of the Tropidurus group east of the Andes (exclud- AxIAL SKELETON ing Uranoscodon), exhibit the loss ofthe medial centrale. 38. Pubic symphysis, anterior margin: (0) 35. Fourth metacarpal and first phalanx of acute; (1) flattened. In the outgroups and most fourth finger (fig. 17): (0) fourth metacarpal of the Tropidurus group, the anterior margin distinctly shorter than third metacarpal -first of the pubis is acute, but in Uracentron and phalanx offourth finger distinctly shorter than Strobilurus the anterior margin is relatively first phalanx of third finger; (1) fourth meta- flat. Possibly this is associated with caudal carpal equal to length of third metacarpal- musculature in these lizards with heavily first phalanx offourth finger distinctly shorter armed tails. than first phalanx of third finger; (2) fourth 39. Anterior caudal vertebrae, neural metacarpal equal to length of third metacar- spines (fig. 18): (0) moderate to high; (1) very pal -first phalanx of fourth finger subequal depressed. Although the neural spines of the to first phalanx ofthird finger. All ofthe Trop- caudal vertebrae are variably developed in idurus group east of the Andes (except Ura- the Tropidurus group, T. bogerti and Tapi- noscodon) exhibit elongation of the fourth nurus have effectively reduced the neural 1992 FROST: TROPIDURUS GROUP OF LIZARDS 23

Fig. 18. Lateral views of third caudal vertebrae, showing development of neural spines. A: Tropidurus hispidus, KU 135268. B: Tropidurus bogerti, RWM 11662. Scales = 2 mm. spines of the anterior caudal vertebrae to a minimum. 40. Caudal vertebrae, autotomy fracture planes (Etheridge, 1967): (0) present; (1) ab- Fig. 19. Lateral view of snouts showing nostril sent. Unique in the Tropidurus group, Ura- morphology and exposure. A: Tropidurus there- centron lacks caudal autotomy fracture planes. sioides, LACM 134136. B: Tropidurus melanopleurus, KU 183472. C: Uracentron azureum, KU NARIAL CHARACTFERS 204989. Nasal scale stippled. Scales = 2 mm. 41. Nostrils (fig. 19): (0) exposed posterolaterally and key-hole shaped or some other 24 AMERICAN MUSEUM NOVITATES NO. 3033

tion into transformation series. For example, most have faint to bold black transverse chest bars not found in the outgroups. Evaluation of the presence of this feature is difficult because of various modifications. Presumably this coloration is a synapomorphy of the Tropidurus group, excluding Uranoscodon. Within the Tropidurus occipitalis and T. peruvianus groups ofspecies, longitudinal paravertebral light stripes are common, particularly from the eye to over the shoulder. Similar stripes are present in some species of Leio- cephalus and in the "Stenocercus" group, so the status of this feature is arguable. I have found only one feature of coloration that is amenable to placement in a transformation series. 42. Ventral thigh and preanal pigmented region (Rodrigues, 1987) (fig. 20): (0) thighs without a well-defined ventral pigmented spot; (1) thigh and preanal region with a well- defined yellow or greenish spot; (2) thigh and preanal region with a well-defined brown to Fig. 20. Pigmented thigh patches. Ventral view black spot. Although some members of the of Tropidurus bogerti, RWM 11659. Scale = 10 "Stenocercus" group show fairly extensive mm. "flash" markings on the ventral side of the thighs, these marks are never the well-defined modification thereof; (1) exposed laterally and in and are widely open; (2) directed anteriorly or an- spots seen the Tropidurus group terolaterally and unconstricted. As in the actually more similar to markings found in "Stenocercus" group and some Sceloporus (Phrynosomatidae). None Leiocephalus (as T. well as other iguanians examined), the Tropi- of the T. peruvianus group or occipitalis durus group plesiomorphically has the edges group shows any kind of thigh spotting, nor of the nostril produced posterovertically and does Uranoscodon. In some taxa the well- has a nasal constriction within the nostril that defined thigh spots are yellow or greenish (i.e., gives the nostril a key-hole or oblong shape. Tropidurus nanuzae, T. melanopleurus, T. Also, in outgroups and in most of the Tropi- spinulosus, Plica [with less sharp edges in Pli- P. durus group the nostril is exposed dorsola- ca plica and lumaria], and Strobilurus). In However, in Tropidurus bogerti, T. other taxa (i.e., Tropidurus bogerti, T. coco- terally. robensis, T. erythrocephalus, T. etheridgei, T. melanopleurus, and Tapinurus the nostril is T. T. T. tor- exposed more laterally and is more widely hispidus, insulanus, hygomi, and open, with the normally produced edges of quatus, and Tapinurus) the thigh markings the nostril reduced. In Plica, Strobilurus, and are dark. Additionally, some species (i.e., T. Uracentron the nostrils are directed more an- etheridgei, T. itambere, and some T. coco- terdorsally. I have left this transformation robensis) show an additional melanistic fleck series unordered because, beyond condition on the belly. However, characterization is suf- "0" being plesiomorphic, I have no compel- ficiently difficult that I have considered the ling reason to polarize conditions "1" and transformation as unordered, although the thigh markings are likely homologous regardless of hue. COLORATION HEMIPENES Members of the Tropidurus group vary in Hemipenes in lizards have been poorly ex- coloration in ways that mostly defy transla- amined since the early work by Cope (1897) 1992 FROST: TROPIDURUS GROUP OF LIZARDS 25

(but see Bohme, 1988), but they provide some evidence here. Arnold (1984) noted that the Tropidurus group is characterized by an accessory dorsal hemipenial muscle, found otherwise only in polychrids. I have not evalu- ated this character in all species, but because Arnold (1984) saw it in Uranoscodon superciliosus, Uracentronflaviceps, Strobilurus torquatus, Plica umbra, Tropidurus peruvianus, and T. torquatus, I consider this a synapomorphy of the Tropidurus group. 43. Hemipenes, condition (Bohme, 1988) (fig. 21): (0) no terminal disks on hemipenial lobes; (1) terminal disks present on hemipenial lobes. In outgroups and members of the Tropidurus group east of the Andes the lobes of the hemipenis do not end in a terminal bare disk. With the exception of Tropi- durus koepckeorum, all Tropidurus group west A B of the Andes have distinctive terminil disks Fig. 21. Hemipenes. A: Plica umbra, KU on the lobes of the hemipenis. T. koepckeo- 147947. B: Tropidurus tigris, KU 167352. Both rum has long hemipenial lobes more like east- x 10. ern Tropidurus. 44. Hemipenes, length oflobes (fig. 21): (0) short; (1) long. Leiocephalus has single-head- between the hemipenial lobes. In Leiocepha- ed, single-sulcate hemipenes that are incipi- lus ornamentation extends far down the shaft ently divided. All members of the "Steno- of the hemipenis. Therefore, outgroup com- cercus" group examined have short-lobed, parison is insufficient to determine the po- bisulcate, bilobate hemipenes. Although with larity of this particular transformation. For respect to outgroups, all members of the purposes of this study I regard this transfor- Tropidurus group have elongate hemipenial mation as nonpolarized (i.e., the "ancestor" lobes, those from west of the Andes, exclud- is coded as "unknown"). ing T. koepckeorum, have the shortest hemipenial lobes. Tropidurus koepekeorum and POCKETS AND FOLDS all species ofthe group east ofthe Andes have The nomenclature ofneck and throat folds elongate lobes (although T. koepckeorum is in iguanian lizards is sufficiently chaotic that on the short side of this variation). Because, I will define my terms. Generally speaking, within the Tropidurus group, lobe length and the folds on the necks oflizards are relatively disking appear to be causally related I have in refrained from recognizing any transforma- conservative topographic position; this notion other than a general elongation of the menclature is applicable to all lizard groups. hemipenial lobes that is a likely synapomor- See figure 25 for a generalized lizard and the phy of the Tropidurus group. nomenclature of folds. More specifically: 45. Hemipenes, ornamentation: (0) caly- A. The antehumeralfold extends anteriorly over ces start below crotch between lobes; (1) caly- the shoulder and may be (usually is) confluent with ces start at a level well above the crotch be- the gular fold should it be present. An interesting tween the hemipenial lobes. Within the exception is in Leiocephalus, the "Stenocercus" group, and in many members of the Tropidurus Tropidurus group, as in the "Stenocercus" group. That is, the antehumeral fold, bordered by group, calyces ornament the lobes of the small scales, extends obliquely under the gular fold hemipenes. In the Tropidurusperuvianus and (bordered by large scales) which becomes obsolete T. occipitalis groups, and T. nanuzae, orna- or may extend anterodorsally all the way to the mentation penetrates on the posterior side of oblique neck fold. In other words, this arrange- the shaft of the hemipenis below the crotch ment ofthe lateral edges ofthe gular fold extending 26 AMERICAN MUSEUM NOVITATES NO. 3033

longitudinal neck

A ' 2 B \/ 2 C Fig. 23. Ventral view showing antegular and gular folds. A: Tropidurus torquatus, USNM 207684 (antegular and gular folds medially incomplete). B: Tropidurus spinulosus, USNM Field (NJS) 126026 (antegular complete and closely ap- Standardized nomenclature of folds. proximating gular fold, ventral mite pockets out- Fig. 22. lined by dashed line). C: Uracentronflaviceps, KU 175317 (antegular and gular fold complete). (1) Antegular fold; (2) gular fold. over the antehumeral fold may be a synapomorphy of the Tropidurinae + Leiocephalinae. In Phymaturus and Liolaemus something near this condition obtains, but the antehumeral fold goes T. habelil) and some "Stenocercus" (e.g., "S." var- over the gular fold. This may be correlated with iabilis). It is under this fold (or its lower extent, the fatty pouches on the sides of the neck in that the antegular fold) that mite pockets form in phry- group. nosomatids and in eastern Tropidurus. B. The gularfold extends immediately anterior F. The term antegularfold represents possibly to the insertion of the arm and may be confluent two nonhomologs. Generally speaking, the ante- with the antehumeral fold. This fold is associated gular fold is a transverse continuation ofthe oblique plesiomorphically with a discontinuity in scale size. neck fold. In some species, like T. stolzmanni, I have seen "gular" folds that are not associated supernumerary antegular folds are variably pres- with this discontinuity but I suspect nonhomology ent. When mite pockets form, the fold of the an- of appearance in these cases. Plica umbra has its tegular (continued as the oblique neck fold) forms gular fold interrupted medially. What confuses the a lobe in front of the scaleless pocket. When two issue is the antegular fold being displaced back- mite pockets obtain underneath the antegular fold wards over the topographic position of the gular they are separated by a smaller fold that divides fold. Plica plica, P. lumaria, and Uracentron do the "mite zone." However, in taxa such as Co- have a complete gular fold. To my mind, only phosaurus (Phrynosomatidae) or some members structural gular folds, that is, folds distinguished of the western Tropidurus group the mite pocket by scale discontinuities are easily characterized. can be very shallow and is detectable only on close C. The dorsolateralfold extends longitudinally inspection. Alternatively, in some species, like anteriorly along the sides and over the insertion many Sceloporus (Phrynosomatidae) or Tropidu- of the arm. The dorsolateral fold is usually con- rus koepckeorum (as well as many of the T. pe- fluent with the antehumerail fold, but sometimes ruvianus group), a lateral antegular fold extends continues forward (as a supra-auricular fold) to a anteriorly under the angle of the jaws where they point over the tympanum. In all species with suf- parallel the longitudinal axis. A fold ofsorts some- ficiently small body scales the dorsolateral fold is times connects these in an "H" shape. In Plica and evident for much of the length of the body. T. spinulosus (to a lesser degree), the antegular fold D. The supra-auricularfold is the continuation is displaced backwards to approach or overlap the ofthe dorsolateral fold anterior to the antehumeral gular fold. fold to a position over the tympanum. G. A longitudinal neck fold frequently is con- E. The oblique neck fold is what Fritts (1974) fluent with the postauricular fold. referred to as the neck fold. This is frequently con- H. A postauricularfold is a continuation ofthe fluent with the antegular fold. The oblique neck longitudinal neck fold that crosses the oblique neck fold may be connected to the antehumeral fold by fold. The postauricular fold may be confluent with a longitudinal neck fold. The oblique neck fold in the nuchal fold if the latter is present. Leiocephalus is very similar to that found in the I. The nuchalfold is the fold at the back of the T. occipitalis group (particularly T. bivittatus and cephalic scales so evident in Leiocephalus, many 1992 FROST: TROPIDURUS GROUP OF LIZARDS 27 iguanids, and Uranoscodon. Sometimes the nuchal fold is confluent with the postauncular fold. J. A rictal fold is present in Plica plica, P. lumaria, Uracentron, Strobilurus, T. melanopleurus, Tapinurus, T. spinulosus, and T. bogerti as an "upward" fold connecting the corner of the mouth with the bottom of the ear. Apparently its development is in part correlated with the development of underlying jaw adductor musculature. The following transformation series (46- 53) are based on variation within these folds and pockets. 46. Gular fold (fig. 22, 23): (0) incomplete medially; (1) complete medially, Both Lei- ocephalus and the "Stenocercus" group have gular folds that are incomplete medially. Pli- ca plica, P. lumaria, Uracentron, and Ura- noscodon are the only members ofthe Tropi- durus group with a medially complete gular fold. 47. Antegular fold (fig. 22, 23): (0) absent or weak and variable; (1) present, strong, well anterior to gular fold; (2) present, strong, Fig. 24. Lateral mite pockets. A: Tropidurus closely approximating or overlapping gular koepckeorum, LACM 49102; showing single mite fold. In the outgroups plesiomorphically and pocket. B: Tropidurus nanuzae, USNM 213514; showing mite pockets under oblique neck fold and in many species of the Tropidurus group a antehumeral fold. C: Tropidurus torquatus, USNM medially complete antegular fold is absent. 207684; showing double mite pockets under Plica, Tropidurus stolzmanni, T. spinulosus, oblique neck fold and lateral extent of antegular T. melanopleurus, Uracentron, and Uranos- fold. codon all exhibit a strong antegular fold. More difficult to characterize, all members of the Tropidurus peruvianus group (with the ex- analogous to how size offused "interparietal" ception of T. tigris), and T. bogerti show weak scales is compared with taxa that lack en- "complete" antegular folds that are individ- larged (= fused) "interparietals." The topog- ually variable in their completeness. I have raphy of folds on the sides of the neck (fig. coded only those species with strong ante- 22) indicates expected position of the antegular folds to have the apomorphic condi- gular fold with some accuracy. tion. Although Leiocephalus lack an ante- 48. Antegular-oblique neck fold mite gular fold, in the species ofthe "Stenocercus" pockets, condition (fig. 24): (0) weak single group that have an antegular fold (e.g., "S." mite pocket (T. occipitalis group, Strobilurus humeralis, "S." crassicaudatus) the fold is torquatus); (1) no mite pocket; complex neck placed well anterior to the gular fold. Also, I folding (Tropidurus peruvianus group, T. bo- am unaware ofany iguanian, other than Plica gerti, Uracentron, Tapinurus); (2) a well-de- plica, P. lumaria, P. umbra, and T. spinu- veloped mite pocket in the upper position losus, that has an antegular fold that approx- (see following transformation); (3) ventrolat- imates or overlaps the gular fold. The pos- eral mite pockets (Tropidurus melanopleurus, terior extension ofthe antegular fold over the T. spinulosus, Plica [weak in P. umbra]); (4) topographic position ofthe gular fold in Plica no obvious mite pocket, although weak de- has caused it to be called a gular fold and has pressions are in the ventrolateral side of an- also resulted, in part, in the misidentification tegular fold (Uranoscodon). ofmany Tropidurus spinulosus as Plica plica. The characterization of mite pockets and The "position" of an antegular fold is eval- the recognition oftransformation series with uated in taxa that lack this fold in a manner the recognized interspecific variation is 28 AMERICAN MUSEUM NOVITATES NO. 3033

gular fold on the ventrolateral side may be homologous with the mite pockets of other taxa. Uracentron, Tropidurus bogerti, and Tapinurus lack mite pockets for much the same reason as the T. peruvianus group, elaboration ofcomplex neck folds. Plicaplica also has complex lateral neck folds, but like P. umbra, which does not have complex lateral neck folds, it has modified the position ofthe gular and antegular folds to such a degree that homology ofsome structures is questionable. However, Plica shares with Tropidurus spinulosus and T. melanopleurus the development ofventrolateral mite pockets under the antegular fold. None of these taxa has lateral mite pockets as in other taxa; on topographic grounds these ventrolateral pockets are likely homoplastic with the lateral mite pockets of other taxa. In the Tropidurus torquatus group, development of mite pockets shows considerable variation. In these species two regions of pocket development can be seen behind the Fig. 25. Neck spines. Plicaplica, AMNH 107590. oblique neck fold, generally separated by a secondary vertical fold. In T. etheridgei and T. cocorobensis both of these pockets are astonishingly difficult, particularly because it weakly developed (most weakly in the Ar- is just these structures that are so useful in gentianian populations); these two pockets are species identification. Mite pockets are regions better defined and much deeper in T. hygomi. ofreduced (or absent) squamation in pockets In T. torquatus the upper pocket is well de- of variable depth, usually located under the veloped and the lower is poorly developed. antegular-oblique neck fold. Generally, these This is similar to the condition in T. erythro- pockets are inhabited by dense colonies of cephalus, T. insulanus, T. itambere, T. mon- brightly colored mites. Complicating the is- tanus, T. oreadicus, and T. hispidus in which sues of homology and polarity of transfor- the lower region of reduced scales is largely mation is the uncertainty of topology owing absent (presumably concomitant with en- to the modification between taxa of the ar- largement ofneck scales). The mite pocket is rangement of neck folds. Classes of condi- extremely enlarged dorsoventrally in T. ore- tions exist that can be discussed, but their adicus and T. montanus. phylogenetic relationships remain mysteri- For purposes of this study I divide this ous. variation into characters of unknown rela- West of the Andes, the Tropidurus peru- tionship to each other. Although I suspect vianus group lacks mite pockets, although the that the "ancestral" condition for the Tropi- complex folds on the sides of the neck may durus group was "0", I have coded the "hy- have obscured these pockets. All members of pothetical ancestor" used for rooting as "un- the T. occipitalis group have invaginations known." behind the oblique neck fold that are invested 49. Oblique neck fold mite pocket, con- with reduced scales and are presumably ho- dition (fig. 24): (0) two mite pockets (T. eth- mologous with pockets behind the lateral neck eridgei [weak in some individuals, particu- folds found elsewhere in the Tropidurus group. larly in Argentina], T. cocorobensis, T. A similar condition obtains in Strobilurus. hygomi, T. torquatus [lower reduced]); (1) a Uranoscodon lacks obvious mite pockets, al- single well-developed mite pocket in the up- though two shallow depressions in the ante- per position indicated (T. erythrocephalus, T. 1 9921992FROST: TROPIDURUS GROUP OF LIZARDS 29 hispidus, T. itambere, T. mucujensis, T. nanuzae); (2) single, very enlarged mite pocket (T. montanus and T. oreadicus). This additional transformation allows resolution of homology hypotheses not possible under the previous transformation. See discussion under previous transformation. 50. Mite pocket in antehumeral fold (fig. 24): (0) absent; (1) present laterally; (2) present ventrolaterally in antehumeral-antegular fold. Tropidurus nanuzae exhibits a very deep antehumeral mite pocket, unique among the iguanians examined. A likely nonhomolo- gous condition obtains in Plica plica and P. lumaria in which ventrolateral mite pockets appear under the gular fold. Because in these two species the gular and antehumeral folds are continuous, I have coded these conditions as part of an unordered transformation, although I consider it unlikely on topographic grounds that these are homologous. 51. Tufts of spines on sides of neck (fig. 25): (0) absent; (1) present. This transformation series is more difficult to assess than one would think. At least in Tropidurus spi- nulosis the development of the spines is ontogenetic and has a strong geographic component ofvariation. All species showing some development have spines in the same topographic position relative to lateral neck folds. No development is seen in the outgroups or Tropidurus west of the Andes. Tropidurus torquatus and T. montanus show slightly enlarged scales in the topographic position of the spine tufts, as do T. melanopleurus, Ura- centron flaviceps, and Uranoscodon. Tropi- durus bogerti, T. mucujensis, T. spinulosus, Plica plica, P. lumaria, Strobilurus torquatus, and Tapinurus semitaeniatus develop distinct tufts of spines in older adults. Expres- sion oftufts is apparently strongly influenced by changes in lateral neck scale size. Regard- Fig. 26. Axillary squamation. A: Tropidurus less, species such as members of the T. pe- hygomi. B: Tropidurus hispidus, showing single ax- ruvianus group, which have small lateral neck illary pocket. C: Tropidurus torquatus, showing scales, lack "tufting." Therefore, "tufting" is multiple axillary pockets. Redrawn from Ro- likely apomorphic for the Tropidurus group drigues (1987). Anterior is to the left. east ofthe Andes, excluding Uranoscodon, or some subset thereof. 52. Rictal fold (fig. 22): (0) absent; (1) pres- the corner of the mouth to under the ear; it ent. In Plica plica, P. lumaria, Strobilurus, is not found in the remainder of the Tropi- Tapinurus, Uracentron, Tropidurus bogerti, durus group, or members of the outgroups. T. melanopleurus, and T. spinulosus, a dis- 53. Supra-auricular fold (fig. 22): (0) ab- tinctive "upward-pointing" fold extends from sent or poorly developed; (1) present, well 30 AMERICAN MUSEUM NOVITATES NO. 3033

variably present in the "Stenocercus" group although this is not the plesiomorphic condition in that group. The multiple axillary pockets of Tropidurus torquatus and Tapi- nurus are visible, but weakly developed in T. bogerti and T. mucujensis. Of the taxa considered, only T. erythrocephalus, T. hispidus, and T. insulanus have a single ("keyhole") axillary pocket. Some specimens of T. montanus, however, show single mite pockets and some T. "hispidus" from the Guyana Region have doubled ones, and some specimens of T. torquatus have the folds reduced to a condition almost identical to that in T. montan- --- us. Some individuals of T. itambere (e.g., MCZ 172883) show axillary depressions that approach the condition found in small T. montanus; for this reason T. itambere is coded as "unknown." T. bogerti has granular regions topographically in the position ofthe pockets of other taxa; for this reason I have coded it as apomorphic. 56. Inguinal granular pocket (fig. 27) (Ro- drigues, 1987): (0) absent or represented solely by a nongranular fold; (1) present. All species in the outgroups and within the Tro- pidurus group that have relatively small scales Fig. 27. Groin squamation. A: Tropidurus his- have something of an inguinal fold so care pidus. B: Tropidurus torquatus, showing groin must be taken in the evaluation of this fea- pocket. Redrawn from Rodrigues (1987). Anterior ture. The difference between distinctly prein- is to the left. guinal and inguinal mite pockets (Rodrigues, 1987) is difficult to characterize because of intraspecific and geographic variation. I have developed. Although present in some mem- therefore only coded presence or absence of bers ofthe outgroups (e.g., "Stenocercus"for- the pocket. Because T. bogerti has a granular mosus), a supra-auricular fold (from the top fold in the topographic position of the fem- ofthe ear to confluence with the dorsolateral oral pocket of other species I have coded it fold) is well developed within the Tropidurus as apomorphic. group only within the T. peruvianus group. Weak (but uncoded) development also can SQUAMATION be seen in some specimens of T. bogerti and T. melanopleurus. Nomenclature of scales follows Smith 54. Axillary pocket, presence (fig. 26): (0) (1946). absent; (1) present. See discussion of varia- 57. Rostral scale, height (Etheridge, tion in axillary pockets under the following 1970a): (0) rostral scale 1.5 to 2 x height of Transformation Series. Although there is adjacent supralabials; (1) rostral scale height considerable variation in the form ofaxillary reduced, less than 1.5 height of adjacent su- pockets, I have coded them as putative ho- pralabials. In the Leiocephalinae and "Steno- mologs. cercus" group, the rostral scale is well ele- 55. Axillary pocket, condition (fig. 26) vated. In T. bogerti, Plica, Uracentron, and (Rodrigues, 1987): (0) present, single; (1) Uranoscodon the rostral scale is reduced to multiple, usually 3, sometimes only 2. Ax- nearly the level of the adjacent supralabials. illary pockets are absent in Leiocephalus and Tropidurus melanopleurus and T. spinulosus 1992 FROST: TROPIDURUS GROUP OF LIZARDS 31

(somewhat more frequently and with a geographic component) also show reduction in height of the rostral scale, but because these do not exhibit the same degree of reduction I have coded these species as plesiomorphic. Although Etheridge (1966) noted variation in nasal-rostral contact within the Tropidurus group, except in clear cases of anomalies, all members of the Tropidurus group have the rostral separated from the nasal scale by at least one row of postrostral scales. 58. Mental scale (fig. 28): (0) enlarged, extending posteriorly well beyond level of adjacent infralabials; (1) reduced, not extending posteriorly well beyond level of adjacent infralabials. In Leiocephalus and the "Steno- cercus" group the mental scale is much larger than the adjacent infralabials. This condition obtains in most ofthe Tropidurus group also. However, in Plica and T. spinulosus the mental scale is reduced and does not extend far beyond the level of the anteriormost infralabials. Although the mental scale of Ura- centron appears on casual inspection to be reduced, this perception is due to the great enlargement of the adjacent infralabials and postmentals. 59. Postmental series (Etheridge, 1968, 1970a) (fig. 28): (0) well defined; (1) poorly defined or absent. In Leiocephalus and the "Stenocercus" group a distinct postmental Fig. 28. Ventral views ofchins showing mental series of scales is evident. This is also the (cross-hatched) and postmental scales (stippled). condition in most of the Tropidurus group, A: Tropidurus torquatus, UNSM 207684. B: Plica except for Strobilurus, Plica, Uranoscodon, umbra, KU 126778. Tropidurus spinulosus, and T. melanopleurus in which the postmental series is difficult to discern from adjacent gulars and subinfrala- cercus" group the infralabials are not ex- bials. Ofthis group, T. melanopleurus has the panded. This is also the condition in most most evident postmentals; I have coded this species in the Tropidurus group. Although species as "1" because the postmental series Tropidurus bogerti, T. melanopleurus, Strobi- is reduced and because of the unusual posi- lurus torquatus, and Tapinurus exhibit some tion of these scales; these' mildly enlarged ventral expansion ofthe infralabials with re- scales may not be homologous with the en- spect to the adjacent supralabials, this con- larged postmentals of other species. dition is approached by other Tropidurus (e.g., 60. Infralabial scales, number (fig. 29): (0) T. torquatus, T. hispidus), making character- 6; (1) 8-9. In outgroups and most ofthe Trop- ization ofa transformation series impossible. idurus group there are 6 infralabial scales. In Uracentron carries this expansion to a much the Tropidurus peruvianus group, however, greater degree and has the only condition here there are 8-9. coded as apomorphic. 61. Infralabials, expansion (Etheridge, 62. Lateral gular scales: (0) imbricate pos- 1968) (fig. 29): (0) infralabials not ventrally teriorly; (1) imbricate laterally. In Leiocepha- expanded; (1) infralabials greatly expanded lus and the "Stenocercus" group, the lateral ventrally. In Leiocephalus and the "Steno- gular scales are imbricate posteriorly. Within 32 AMERICAN MUSEUM NOVITATES NO. 3033 superciliaries

C DD infralabials Fig. 29. Scale characters ofthe side ofthe head. A: Tropidurus atacamensis, KU 161982. B: Tropidu- rus hispidus, KU 127227. C: Plica umbra, KU 126778. D: Uracentronflaviceps, KU 126781. Infralabials, superciliaries, and preocular-subocular series shaded on each view, although only labeled once. the Tropidurus group this is also true in Ur- the extreme condition and additionally have acentron and T. bogerti. All other species of the direction ofimbrication ofthe head shields' the Tropidurus group have the scales imbri- reversed over the entire head. cate posterolaterally. In some species of the Possibly this imbrication would increase T. peruvianus group the lateral gulars are the handling time ofpredatory snakes, which granular but still show evidence oflateral ori- are known to take cues from the direction of entation. scale imbrication of prey items (Greene, 63. Scales of frontonasal region (fig. 25): 1976). (0) imbricate posteriorly or no imbrication 64. Superciliary scales (Etheridge, 1 970a) evident; (1) weakly imbricate anteriorly; (2) (fig. 29): (0) not or only weakly produced ver- all head shields strongly imbricate anteriorly. tically to form a longitudinal crest; (1) pro- Unique within the iguanians examined, duced vertically to form a longitudinal crest. Tropidurus spinulosus, T. melanopleurus, The superciliaries of Plica and Uranoscodon Strobilurus, Plica, Tapinurus, and Uracen- are produced vertically conspicuously more tron have the scales ofthe frontonasal region than in other members of the Tropidurus imbricate in an anterior direction. In some group or outgroups. individuals of Plica umbra and Uracentron, 65. Circumorbital series (fig. 30): (0) in one the anterior direction of imbrication is only row between the supraoculars and the me- detectable by lifting the stratum corneum with dian head shield; (1) in two rows between the a probe. Tropidurus bogerti is coded as "un- supraoculars and the median head shield. In known" because some individuals appear to most species of the Tropidurus group as well have some subtle anterior imbrication and as Leiocephalus and relevant members ofthe others do not. Plicaplica and P. lumaria show "Stenocercus" group there is only one row of 1992 FROST: TROPIDURUS GROUP OF LIZARDS 33 circumorbitals separating the supraoculars from the median head shields. However, in Tropidurus melanopleurus, T. spinulosus, Plica, Strobilurus, and Uracentron there are two distinct rows or circumorbitals, at least posteriorly. Because a circumorbital series is autapomorphically indistinguishable from the supraoculars in Uranoscodon this species was coded as "unknown" in the analysis. 66. Circumorbitals (fig. 30): (0) small; (1) enlarged at the expense of the supraoculars. In Uracentron the circumorbitals, normally small in the other members ofthe Tropidurus group and outgroups, are enlarged at the expense of the supraocular scales. 67. Interparietal (Smith, 1946; Etheridge and de Queiroz, 1988) (fig. 33): (0) not enlarged, smaller than interorbital distance; (1) enlarged, larger than interorbital distance. With the exception of the Phrynosomatidae, the Tropidurus group is unique in the pos- session of an "enlarged" interparietal. "En- largement" is not really an appropriate term because the interparietal of the Tropidurus group is clearly a sutured aggregation of parietal scales as in other iguanians. In Ura- noscodon the individuality of the scales is evident because of their separate elevation regardless of their edge-to-edge suturing. In most specimens ofwestern Tropidurus, there is generally some evidence ofincomplete suturing around the periphery of the interparietal scale (fig. 30). Etheridge (1970a: 242) noted that Uranoscodon has a relatively small interparietal, but because it does not obvi- ously have the "enlarged" interparietal com- of in posed fewer subsidiary scales than other Fig. 30. Dorsal head squamation; circumor- members ofthe Tropidurus group, I have not bitals and interparietals shaded. A: Tropidurus used this difference in my analysis. koepckeorum, LACM 49102 (frontonasal scales 68. Interparietal length (Etheridge, 1968) pavemented). B: Tropidurus spinulosus, LACM (fig. 30): (0) subequal to significantly less than 126318 (frontonasal scale imbricated anteriorly). width; (1) significantly longer than wide. In C: Uracentronflaviceps, KU 175318 (frontonasal order to evaluate the polarity of this feature scales imbricated anteriorly). I circumscribed the "parietal scales" in the outgroups so as to estimate the dimensions ofthe "enlarged" interparietal (see discussion (0) 0-1; (1) 2 or more. In both Leiocephalus in previous transformation series). Uracen- and the "Sternocercus" group, as well as most tron and T. itambere are unique within the of the Tropidurus group, there is no more Tropidurus group in having an interparietal than one row ofloreolabials between the sub- substantially longer than wide. ocular and the supralabials. In Plica there are 69. Rows of scales between subocular and at least two rows of loreolabials penetrating supralabials (Etheridge, 1970a [part]) (fig. 29): between the subocular and supralabials. In 34 AMERICAN MUSEUM NOVITATES NO. 3033

-_*- anterior condition obtains in most of the Tropidurus group. However, in Plica, Strobilurus, Ura- centron, and Uranoscodon the orbit is underlain by at least three scales caused by fragmentation of the preocular-subocular series. Uranoscodon and Plica umbra have continued this fragmentation to the point that the preocular-subocular series is obscure. 71. Preauricular fringe (fig. 31): (0) present, ear canal deep, a continuous fringe of scales partially to nearly completely covering ear opening; (1) reduced, ear canal deep; (2) auricular scales reduced, ear canal deep, a lower lobule with several short spines present; (3) auricular fringe absent, the ear canal shallow. In relevant members of the out- /W groups and most ofthe Tropidurus group, the external ear canal is partially covered by a fringe of scales. In the T. peruvianus group the fringe is reduced through "granularization" concomitantly with the granularization ofbody scales. In some species (i.e., Tropidu- C~~ rus spinulosus, T. bogerti, T. melanopleurus, '-- Plica plica, P. lumaria, Strobilurus, and Tap- D inurus) the fringe is reduced, and a short lob- Fig. 31. Auricular fringing and lobing. A: ule on the ventral limit ofthe anterior margin Tropidurus koepckeorum, LACM 49102. B: Tropi- of the ear canal is present. Tapinurus semi- durus atacamensis, KU 161980. C: Tropidurus taeniatus is intermediate between "0" and spinulosus, USNM Field 99128. D: Uranoscodon "2" but was coded as "2", because it exhibits superciliosus, KU 127230. the fleshy lobe. Plica umbra, Uranoscodon, and Uracentron have a more reduced con- occasional specimens (uncoded) of Tropidu- dition; the entire fringe or lobule is absent. rus spinulosus more than one row of scales Because the "reduced" condition (2) involves may penetrate between the supralabials and the elaboration ofa fleshy lobule also not seen subocular. Uranoscodon may appear, on cur- in condition "3", it does not follow neces- sory examination, to have multiple rows of sarily that absence (3) must be derived from scales below the subocular. This illusion is condition "1" rather than "0" or "2". I have caused by uncertainty as to what constitutes therefore regarded these conditions as an a subocular, because this species has a highly unordered set, although the "ancestral" con- fragmented subocular series. However, oc- dition is clearly "0". casional specimens show enough enlarge- 72. Middorsal scale row (Etheridge, 1966 ment ofthe subocular series so that it can be [part]): (0) present; (1) absent. I have not taken seen that only one scale row penetrates be- degree of development (i.e., scale enlarge- tween this row and the supralabials. ment or elevation) into consideration in this 70. Subocular (Etheridge, 1 970a) (fig. 29): transformation series because variation in (0) entire-0-1 preoculars; (1) divided-at development is continuous and exceedingly least 2 preoculars in contact with the orbit; difficult to characterize. In Leiocephalus (ex- (2) subocular-preocular series so fragmented cept L. pratensis) and plesiomorphically in as to be obscure. In the "Stenocercus" group the "Stenocercus" group (as in all iguanians) and Leiocephalus, the orbit is underlain by the middorsal scale row is present and en- an elongate subocular and (variably) by a sin- larged. In the T. torquatus group, Tapinurus, gle preocular in the same series. Also, this and Uracentron the middorsal scale row is 1992 FROST: TROPIDURUS GROUP OF LIZARDS 35 not identifiable. In the T. heterolepis subgroup (T. atacamensis, T. heterolepis, T. B C quadrivittatus, and T. theresiae) ofthe T. peruvianus group, the middorsal scale row is usually identifiable on the neck (though not enlarged) but is usually unidentifiable along the back. 73. Paravertebral scales (Boulenger, 1885): (0) keeled; (1) not keeled. In the outgroups and most of the Tropidurus group, the paravertebral scales are imbricate and keeled. In the Tropidurus peruvianus group, Tapinurus, Tropidurus melanopleurus, and Uracentron azureum these scales are extremely weakly keeled or unkeeled. 74. Lateral body scales: (0) imbricate, keeled; (1) granular and juxtaposed. In both outgroups and most ofthe Tropidurus group, the lateral body scales are imbricate and mucronate. In the Tropidurus peruvianus group, though, the lateral body scales are granular ("pebbly") and juxtaposed. Tapinurus approaches this condition, but is not "pebbly" as in the Tropidurus peruvianus group. 75. Caudal scales (fig. 32): (0) tail un- armed, longer than head + body; (1) tail armed with heavy mucrons, roughly terete and subequal to length of head + body; (2) tail armed with heavy mucrons, dorsoventrally flattened and shorter than head + body. Fig. 32. Caudal scales in dorsal view. A: Tropi- In Leiocephalus and plesiomorphically with- durus etheridgei, KU 73442 (SVL = 65, tail = 91). in the "Stenocercus" group the caudal scales B: Strobilurus torquatus, MCZ 154241 (SVL = 45, are only weakly mucronate. In Strobilurus, tail = 40). C: Uracentron azureum, KU 204988 and even more so in Uracentron, the caudal (SVL = 36, tail = 19). Scales = 10 mm. scales are so strongly mucronate that their tails should be considered armed and dan- RESULTS AND DISCUSSION OF gerous. POSSIBLE ERRORS 76. Ventral scales: (0) smooth; (1) keeled. With the exception ofthe few derived species RESULTS of "Ophryoessoides," Proctotretus azureus, For the characters under analysis Tropidu- and Leiocephalus herminieri the keeled ven- rus heterolepis, T. quadrivittatus, and T. ther- tral scales of Uranoscodon superciliosus and esiaewere identical to T. atacamensis. Tropi- Plica umbra are unique among the Tropidu- durus thoracicus, T. theresioides, T. tigris, and rus group and the immediate outgroups. T. yanezi were identical to T. peruvianus. 77. Upper thigh scales: (0) not heavily mu- Tropidurus habelii was equivalent to T. biv- cronate; (1) heavily mucronate. Strobilurus, ittatus, Tropidurus etheridgei equivalent to T. unique within the Tropidurus group and de- hygomi, and Tapinurus pinima equivalent to rived with respect to the outgroups, has T. semitaeniatus. Plica lumaria is identical heavily armed thigh scales. Tropidurus itam- to Plica plica for characters under discussion. bere also has strongly mucronate scales on Tropidurus cocorobensis as characterized in the legs, but nothing approaching the con- the data matrix differs from T. hygomi solely dition in Strobilurus. by "unknown" assignments. 36 AMERICAN MUSEUM NOVITATES NO. 3033

Fig. 33. Strict consensus tree, and one of the 36 equally parsimonious trees generated for the Trop- idurus group data. Length = 169, ci = 0.568. Numbers on stems are noted in Appendices 3 and 4 and discussed under Results.

The reduced data matrix was subjected to ported trees discovered. The "regions" of analysis as described under "Methods." One supported alternative topology are depicted tree was discovered (fig. 33) that was parsi- in figure 34. monious under both accelerated and delayed The following discussion refers to figure 33. transformation. Thirty-six equally parsimo- Stem 1 (the Tropidurus group) is well sup- nious trees (length = 169, ci = 0.568; length ported by five unreversed transformations: = 157, ci = 0.535, excluding a priori auta- 1 1.1 (general reduction of bone mass result- pomorphies and group synapomorphies) were ing, among other things, in the reduction of discovered that had some justification under the quadrate process ofthe squamosal as well either accelerated or delayed character opti- as the enlargement of the superior fossa of mization. Another 16 trees could have been the quadrate), 12.1 (mild reduction in the counted in this number but they depended definition of the alveolar shelf of the man- on "justification" from arbitrary assignment dible [possibly related to transformation 1 1]), of an "unknown" character. The number of 27.1 (enlargement of the sternum), 44.1 unrejected trees, i.e., those whose topologies (elongation of the hemipenes), and 67.1 (fu- were logically consistent with the data but sion ofparietal scales to form a large, distinct whose topologies went beyond data support interparietal scale). Additionally, this stem is numbered 26,588, this dependent on 6 supported by 28.1 (a shortening of the inter- "regions" in the cladogram ofalternative but clavicle median process), which is reversed unsupported arrangements. Of the 36 sup- in Tropidurus bogerti, T. spinulosus, and Ura- ported trees only one was supported unam- centron azureum, and 62.1 (lateral imbrica- biguously (i.e., not dependent on method of tion of the throat scales), which is reversed character optimization). This tree (fig. 33) is in T. bogerti and Stem 19 (Uracentron). This the same as the strict (Sokal and Rohlf, 198 1) is a well-corroborated group. and Adams (1972) consensus of the 36 sup- Uranoscodon superciliosus is supported as 1992 FROST: TROPIDURUS GROUP OF LIZARDS 37

Fig. 34. Supported alternatives for various parts of the cladogram. the sister taxon of the rest of the Tropidurus Stem 3 (other species ofwestern Tropidurus). group (Stem 2). Autapomorphies are numer- T. koepekeorum is not characterized by any ous (17), although only one (37.1), fringed apomorphies here analyzed but some of its toes, seems to be unique to this taxon. A features are likely apomorphic (see below). surprising number ofshared homoplasies with Stem 3, subtending the members ofthe Trop- Plica umbra (20) as well as other species in idurus occipitalis and T. peruvianus groups of its "neighborhood" make it understandable Dixon and Wright (1975) is corroborated by that these taxa have been considered closely a single, but striking feature, hemipenis with related by many authors. Stem 2, the sister terminal disks (43.1). There is currently no taxon of Uranoscodon is corroborated by only evidence for the monophyly of the T. occip- two synapomorphies, although these are italis group, even excluding the former mem- unique and unreversed and easily character- ber, T. koepekeorum. The T. peruvianus group ized: 13.1 (development ofa lingual coronoid (Stem 4), however, is well corroborated by process of the dentary overlapping the an- three unique, unreversed synapomorphies: terior lingual leg of the coronoid), and 33.1 53.1 (supra-auricular fold), 60.1 (8-9 infra- (elevation of the head of the humerus). To labial scales), and 74.1 (granular lateral body this meager list can be added one other fea- scales). Additionally, 48.1 (complex neck ture, black transverse bars across chest and folding), also known homoplastically in T. upper arms. This feature is difficult to char- bogerti, Tapinurus, and Uracentron, 71.1 (re- acterize a priori and is obscured in numerous duced preauricular fringing [presumably re- species by increasing black and other ob- lated to reduction of body scales in general], scuring patterns. However, Uranoscodon found in a number of other species homo- clearly does not have anything like transverse plastically, and 73.1 (reduced scale keeling), black bars on the chest and all of the re- also found homoplastically in Tapinurus, maining species patristically near the "base" Tropidurus melanopleurus, and Uracentron of Tropidurus do, both east and west of the azureum, support this clade. The Tropidurus Andes. heterolepis subgroup (T. atacamensis, T. het- Stem 2 subtends a trichotomy, although erolepis, T. quadrivittatus, and T. theresiae) under some optimizations Tropidurus ofthe T. peruvianus group has its monophyly koepckeorum is placed as the sister taxon of supported by 72.1 (loss ofmiddorsal enlarged 38 AMERICAN MUSEUM NOVITATES NO. 3033

scale row), also seen in the former Tropidurus Stem 6 has no unique unreversed apo- torquatus group and Uracentron. The mono- morphies but is supported by 25.1 (clavicle phyly of this group is not particularly sur- not flanged), reversed in Plica umbra and prising inasmuch as all species seem to be Uracentron (Stem 19), 45.1 (ornamentation predominantly intertidal feeders (Dixon and of hemipenes starts above crotch between Wright, 1975; Ortiz-Zapata, 1980a). lobes), also in Uranoscodon, and 72.1 (loss Stem 5 (Tropidurus and generic satellites of middorsal enlarged scale row), apparently east of the Andes) is highly corroborated by regained at Stem 15, then lost again in Ura- nine unique unreversed synapomorphies: 1.1 centron (Stem 19) and also appearing in the (increased skull size), 7.1 (conspicuously en- Tropidurus heterolepis subgroup. This feature larged nutritive foramina of maxilla), 12.2 has traditionally been the sole justification of (alveolar shelf of mandible strongly eroded), the "Tropidurus torquatus group," which at 14.1 (long posterior extension of dentary), this point has little other support. 16.1 (angular reduced), 20.1 (enlargement of Stem 7 is united by only two features, 49.1 anterior maxillary teeth), 33.2 (ball-like head (single large mite pocket on neck) and 54.1 of humerus), 34.1 (loss of medial centrale), (axillary pocket present), both with homo- and 35. 1 (elongation of fourth metacarpal). plasy; 49 is modified frequently elsewhere and To the list can be added: 15.1 (anterior sur- 54 is reversed on Stem 15. Axillary pockets angular foramen captured by contact of cor- are extremely difficult to characterize for phy- onoid and surangular), reversed in T. bogerti, logenetic analysis and this stem is very poorly 17.2 (posterior mylohyoid foramen between corroborated. The same is true of Stem 8, splenial and dentary), reversed in T. spinu- supporting the monophyly ofT. itambere and losus and Uracentron (Stem 19), 30.1 (supra- T. erythrocephalus which is supported by the scapular fenestrations), reversed in Uracen- homoplastic feature 56.1 (groin granular tron (Stem 19), and 48.2 (lateral mite pocket), pocket) which appears elsewhere in the for- reduced and modified a number of places mer T. torquatus group. I consider this as- above this level in the cladogram. Addition- sociation, along with most other phylogenetic ally, a synapomorphy of this clade is pig- structure within the former T. torquatus group mented thigh patches. This is, after Stem 1, to be poorly corroborated. That T. itambere the most highly corroborated stem in the resembles T. etheridgeiin aspects ofform and analysis. coloration that are difficult to characterize for Tropidurus nanuzae has two unreversed phylogenetic analyses only strengthens this synapomorphies: 26.1 (loss of sternal fonta- suspicion. nelle) and 50.1 (mite pocket in antehumeral Stem 9 is supported solely by rib formula fold). A third feature, 22.1 (posterior max- (31.1). Although Etheridge (1962, 1964) has illary teeth appear brachydont from the labial documented that this can be astonishingly side) is also shared with Uracentronflaviceps, variable in Sator and other phrynosomatids, Plica, and Uranoscodon superciliosus. I have modalities can be established that seem to be been unable to see T. amathites or T. divar- informative. Nevertheless, the sample sizes icatus, both species considered close to T. examined here were small and, although I nanuzae by Rodrigues (1986) on the basis of found no intraspecific variation, it is con- the loss ofthe sternal fontanelles. Apparently, ceivable that taxa have been mischaracter- T. amathites has antehumeral mite pockets ized. and T. divaricatus lacks them. Also, the three Stem 10 is justified by the single character species in the T. nanuzae group ofRodrigues 55.1 (double axillary mite pocket). This fea- (1986) share a presumptively apomorphic ture is reversed on Stem 15 concomitant with karyotype (Kasahara et al., 1987). As it stands, the reversal in Transformation 54 to 54.0 T. nanuzae does not "fit" well anywhere, and (axillary pocket absent). This character is pe- sufficient homoplasy shared with the Plica- culiar and surprisingly difficult to character- Uracentron-Strobilurus region of the clado- ize. That some specimens of T. "hispidus" gram may suggest that I have misplaced it from the Guyana Region have double axil- badly (see comment below). lary pockets and that some T. montanus are 1992 FROST: TROPIDURUS GROUP OF LIZARDS 39 annectent between the 1- and 2-pocket con- manni, 54.0 (loss of axillary pocket), 56.0 dition make this a difficult transformation. (loss ofgroin pocket), 59.1 (postmental series Stem 1 1, like Stem 10, is poorly corroborat- reduced), reversed in Uracentron (Stem 19), ed, with only one homoplastic character jus- and 72.1 (middorsal scale row present). tifying it: 56.1 (groin granular pocket), this Stem 16 has a number ofsynapomorphies: feature being found also on Stem 8, leading 5.1 (nasal spine ofpremaxilla broad), also in to T. itambere and T. erythrocephalus. Stem Uranoscodon, 8.1 (maxillopalatine foramen 12 is supported by scale "tufting" on the sides very large), 21.1 (flaring posterior maxillary ofthe neck. This is lost in Plica umbra, Ura- teeth), also in Tapinurus and T. hispidus, and centron azureum, and apparently similar but 23.1 (posterior maxillary teeth set vertically), weakly developed squamation can be seen in also in T. nanuzae, 29.0 (scapular deflection Uranoscodon superciliosus. reversed), 41.2 (nostrils directed anterolat- Stem 13 is well corroborated by two un- erally or anteriorly and unconstricted), and reversed unique apomorphies: 35.2 (fourth 70.1 (subocular scale divided), also in Ura- metacarpal lengthened) and 63.1 (scales of noscodon. frontonasal region imbricate anteriorly), al- Plica (Stem 17) is supported by only two though T. bogerti was coded "unknown" for nonhomoplastic characters, 10.1 (squamosal this feature because ofcharacterization prob- bone curved around posterior end of tem- lems. Additionally, several other homoplas- poral fenestra) and 69.1 (two or more rows tic features support this stem's reality: 29.1 of scales between the subocular and the su- (scapular deflection with concomitant reduc- pralabials). Other features that support the tion of the scapular fenestra of the scapulo- monophyly of this group are: 22.1 (posterior coracoid), reversed on Stem 16, 52.1 (pres- teeth appear brachydont when viewed from ence ofa rictal fold), reversed in Plica umbra, the labial side), also in T. nanuzae and Ura- and 71.2 (auricular scales reduced), this being centron flaviceps, 47.2 (antegular fold over- further modified in Plica umbra and Uracen- laps gular fold), also in T. spinulosus, 58.1 tron (Stem 19). Stem 14, uniting T. bogerti (mental scale reduced), also in T. spinulosus, with Tapinurus, is supported by two non- and 64.1 (superciliary scales produced ver- homoplastic features associated with being tically to form a crest), also in Uranoscodon. flat: 6.1 (nasal bones reduced) and 39.1 (de- The members of the Plica plica group (P. pressed neural spines oftail vertebrae) as well plica and P. lumaria) are otherwise extremely as 24.1 (loss of pterygoid teeth), which is different from Plica umbra in almost all other highly homoplastic. Tapinurus is a highly ways. Plica lumaria and P. plica are associ- apomorphic group of three species of which ated by all features that are treated as auta- the following are synapomorphies rather than pomorphies of Plica plica in this analysis. autapomorphies of T. semitaeniatus: 3.1 Like the three species ofPlica, Strobilurus (skull compressed), 6.2 (nasal bones very re- and Uracentron have long been associated duced), 18.1 (posterior position of posterior with each other, although the resemblances mylohyoid foramen), also in Uracentron when enumerated are not overwhelming; al- (Stem 19), 19.1 (premaxillary teeth 4-5), also most as much associates Plica umbra with homoplastic in alternative placement else- Uracentron as does Strobilurus with Uracen- where, 21.1 (flaring maxillary teeth), also in tron (i.e., a one-step difference in total tree T. hispidus and at Stem 16, 29.2 (strongly length). However, the weight ofthe evidence flexed scapulocoracoid), 31.0 (reversal to 3 does support a special relationship between sternal ribs), and 32.1 (long xiphisternal rods Strobilurus and Uracentron. This associative associated with the pectoral musculature). Stem 18 is supported by two synapomor- Stem 15 is well corroborated, but only one phies: 38.1 (anterior margin of pubis not of its synapomorphies is unique and unre- acute) and 75.1 (armed caudal scales). The versed: 65.1 (circumorbital scales in two rows). tail structure is otherwise quite dissimilar; Others include 42.1 (ventral thigh region with Strobilurus has a terete tail as is found in such yellow patches), also in T. nanuzae, 47.1 species as Stenocercus roseiventris and Ura- (complete antegular fold), also in T. stolz- centron has a spatulate tail similar in some 40 AMERICAN MUSEUM NOVITATES NO. 3033

Fig. 35. Alternative tree, length = 171. ways to that in Hoplocercus (Hoplocercidae). a cladogram of length 171 (two steps longer Uracentron monophyly is not problematical, than the 169-step preferred cladogram). This supported by four unique apomorphies and allows monophyly of the Tropidurus torqua- a plethora of others: 17.0 (posterior mylo- tus group (including Tapinurus); that is, the hyoid foramen between dentary and angular), traditional synapomorphy ofthe T. torquatus also in T. spinulosus, 18.1 (posterior mylo- group + Tapinurus, loss of middorsal scale hyoid foramen placed far back on mandible), row, is judged unreversed. The enlarged me- also in Tapinurus, 25.0 (clavicle strongly dial dorsal scale row of T. melanopleurus, T. flared), also in Plica umbra, and plesiomor- spinulosus, Plica, and Strobilurus would be phy below Stem 6, 30.0 (reversal to no su- judged plesiomorphic rather than derived prascapular fenestrae), 36.1 (claw of first toe from the T. torquatus condition. Also, re- strongly flexed), 40.1 (no fracture planes in versals having to do with axillary and groin caudal vertebrae), 42.0 (thighs without ven- granular pockets would disappear and both tral pigmented patches), 46.1 (gular fold com- groups ofthigh colors (yellow vs. dark brown plete medially), also in Plica and Uranosco- or black) would be historically connected don, 59.0 (reversal to postmental series rather than have the yellow hue appear in- enlarged), 61.1 (infralabials greatly expand- dependently in T. nanuzae and in the T. mel- ed), 62.0 (lateral gular scales imbricated pos- anopleurus-Uracentron group. As evidenced teriorly), also in T. bogerti, 66.1 (circumor- by the increased tree length, however, the cost bitals enlarged), 68.1 (interparietal much is a reduction ofparsimony, showing itselfin longer than wide), also in T. itambere, 71.3 additional homoplasy in rib formula (addi- (auricular fringe absent, ear canal shallow), tional change from 3 to 4 sternal ribs in the 72.1 (middorsal scale row absent), also in the stem leading to T. melanopleurus, T. spinu- former Tropidurus torquatus group, and 75.2 losus, Plica, Strobilurus, and Uracentron, as (spatulate tail). well as within the T. torquatus group); scapulocoracoid flexing would have to occur in- COMMENT ON THE TROPIDURUS TORQUATUS dependently in the T. melanopleurus-T. spi- GROUP AND ON STABILITY OF nulosus clade as well as in the T. bogerti- THE CLADOGRAM Tapinurus clade (both of which are notably saxicolous clades). The elongation of the That the most parsimonious cladogram (fig. fourth metacarpal would have to occur twice: 33) is unstable is obvious. Figure 35 shows in the T. bogerti-Tapinurus clade and inde- 1992 FROST: TROPIDURUS GROUP OF LIZARDS 41

Strobilurus Strobilurus Western Tropidurus Pica U\rcentron Uracentron Tapinurus Tropidurus" Tapinurus Plica "Westem )don "Eastern Tropidurus" Uranoscodon "Easter Tropidurus"

Etheridge and This study de Queiroz (1988)

Fig. 36. Tropidurus group tree of Etheridge and de Queiroz (1988) and of the present study. pendently in the T. melanopleurus-Uracen- relationship of Plica with Strobilurus and tron clade. Neck spine tufting and the ap- Uracentron. They disagree, however, in the pearance of the rictal fold (associated with placement of this combined group and the underlying muscle development) would also western Tropidurus group. Etheridge and de have increased homoplasy. So, even if rib Queiroz (1988) thought the Plica- Uracen- formula (Transformation 31), scapulocora- tron-Strobilurus group to be the sister taxon coid flexion (Transformation 29), elongation of Tropidurus + Tapinurus. This is based on ofthe fourth metacarpal (Transformation 35), the view that a medially incomplete gular fold rictal fold (Transformation 52), and neck scale is a synapomorphy of Tropidurus + Tapi- tufting (Transformation 51) are excluded from nurus. However, a medially incomplete gular the analysis, this alternative tree is still only fold is a synapomorphy of the Tropiduridae marginally more parsimonious (i.e., two steps and is a plesiomorphy at this level of uni- shorter) than the preferred cladogram, and versality (Frost and Etheridge, 1989). Once would still leave Tropidurus paraphyletic, this problem is removed the only point of even though a modified T. torquatus group logical inconsistency is in the placement of (including Tapinurus) would survive. And, the western Tropidurus group, which Ether- inasmuch as the most parsimonious clado- idge and de Queiroz (1988) derived from the gram correlates well with trends in overall eastern Tropidurus group. None of the fea- similarity and longer trees do not, there is tures documented in my study that place little that should drive us to prefer a longer western Tropidurus outside a group com- tree than we are required to. Regardless of posed of eastern Tropidurus, Plica, Uracen- the choice of overall cladograms, the phylo- tron, Strobilurus, and Tapinurus were avail- genetic structure among species within the able to these authors, so, with the exception former Tropidurus torquatus group is argu- ofthe gular fold character, their evidence can able, with considerable homoplasy, and is so be viewed as less complete rather than in unstable as to not promote much confidence conflict with that presented here. in its accuracy. CONCLUSION COMPARISON WITH THE HYPOTHESIS OF This analysis has forced the data to yield ETHERIDGE AND DE QUEIROZ (1988) only some of its historical signal. Well-cor- The only previous hypothesis of relation- roborated monophyletic groups are: (1) the ship within the Tropidurus group is that pro- Tropidurus group; (2) western Tropidurus, exposed by Etheridge and de Queiroz (1988). cluding T. koepekeorum; (3) the T. peruvi- Their tree and a diagrammatic rendition of anus group; (4) the T. heterolepis subgroup my most parsimonious tree are shown in fig- ofthe T. peruvianus group; (5) the taxon sub- ure 36. The two trees concur in the placement tended by Stem 5 in figure 33 (the traditional of Uranoscodon as the sister taxon of the re- T. torquatus group, T. melanopleurus, T. spi- maining Tropidurus group as well as in the nulosus, Tapinurus, Plica, Strobilurus, and 42 AMERICAN MUSEUM NOVITATES NO. 3033

Uracentron); (5) T. bogerti + Tapinurus; (6) Gauthier, 1986), defined solely by plesio- T. melanopleurus + T. spinulosus + Plica + morphy with respect to its presumptive sister Strobilurus + Uracentron; (7) Plica + Strobi- taxon, Tropidurus. The problem with this ap- lurus + Uracentron; (9) Uracentron. Evi- proach is that it does not invite additional dence for or against a monophyletic T. tor- evaluation of the proposition of monophyly quatus group (including Tapinurus) is of western Microlophus + T. koepekeorum equivocal without resorting to adaptation ar- and is merely an easy bookkeeping conven- guments. Other supported but still arguable tion. The advantage ofpartitioning Tropidu- relationships are Plica and Strobilurus + rus is that the stem supporting Tropidurus Uracentron. east of the Andes (Stem 5) is highly corroborated and without partitioning would be overlooked. At this time I think it prudent TAXONOMY PROPOSED to take the second, four-genus alternative: It is clear that the current generic taxonomy (1) Uranoscodon: a monotypic genus with of the Tropidurus group is not logically con- at least one unique autapomorphy and the sistent (Hull, 1964; Wiley, 1981) with the sister taxon of the remaining Tropidurus hypothesized consensus phylogeny of the group. group presented (fig. 33), or with any of the (2) Plesiomicrolophus: a monotypic genus most parsimonious trees discovered (or even for Tropidurus koepekeorum. with trees that are not particularly parsimo- (3) Microlophus: a genus for all species of nious, e.g., fig. 35). In designing a generic former western Tropidurus, excluding Ple- taxonomy of the Tropidurus group, I have siomicrolophus koepekeorum, diagnosed by been guided by two constraints: (1) The tax- having disked hemipenes. onomy adopted must be logically consistent (4) Tropidurus: a genus for all former with the recovered phylogenetic pattern (i.e., Tropidurus east of the Andes as well as the the taxonomy must not mislead about re- species formerly in Tapinurus, Plica, Ura- covered phylogenetic history); (2) The tax- centron, and Strobilurus. onomy should be minimally perturbable, that Additionally, I propose that the Tropidurus is, questionable stems should, as much as group be recognized formally as a tribe, Trop- possible, remain unnamed. I have not felt idurini, the sister taxon of a new tribe Steno- obliged to name all suprageneric or subge- cercini (the former Stenocercus group) within neric taxa. There are two attractive alterna- the Tropidurinae of the Tropiduridae (Frost tives for the taxonomy to be selected: and Etheridge, 1989). See Taxonomic Ac- (1) Two genera: Uranoscodon and a single counts (below) for diagnoses of these taxa. genus, Tropidurus, for the group now composed of Tropidurus, Tapinurus, Uracentron, TAXONOMIC ACCOUNTS Plica, and Strobilurus. The advantages ofthis AND DIAGNOSES arrangement are that only nine species have The taxonomic accounts are designed to be their generic names changed, although these nested within those provided by Frost and nine species are highly apomorphic and dis- Etheridge (1989) for taxa ofmore general uni- similar from the traditional eidos of Tropidu- versality. The diagnostic features listed below rus. mention "useful" characteristics, regardless (2) Four genera: Uranoscodon, Microlo- of level of universality; apomorphies are in phus for species offormer Tropidurus having bold type. The Tropidurinae account is in- disked hemipenes, a new genus for Tropidu- cluded only as a referent collective for Ster- rus koepekeorum, and Tropidurus for species nocercini and Tropidurini. east of the Andes (including former Tapinu- rus, Plica, Uracentron, and Strobilurus), oth- TROPIDURINAE BELL, 1843 er than Uranoscodon. One might argue that Tropiduridae Bell, 1843: 1. Type genus: Tropidu- because T. koepekeorum is the likely sister rus Wied-Neuwied, 1825. See comment under taxon ofother species ofwestern Tropidurus, Tropidurinae. it would be more prudent to place it in a Ptychosauri Fitzinger, 1843: 16. Type genus: Pty- collective, Microlophus* (a metataxon sensu chosaurus Fitzinger, 1843 (= Plica Gray, 1831). 1 9921992FROST: TROPIDURUS GROUP OF LIZARDS 43

Steirolepides Fitzinger, 1843: 17. Type genus: DISTIuBUTION: Western South America Steirolepis Fitzinger, 1843 (= Tropidurus Wied- from northern Colombia and coastal Ecuador Neuwied, 1825). to Bolivia, coastal Argentina, and the Upper ?Heterotropides Fitzinger, 1843: 17. Type genus: Amazon Basin of Brazil; coastal southern Heterotropis Fitzinger, 1843 (a nomen dubium) Brazil and Uruguay (fig. 37). (= Ophryoessoides Dumeril and Dumeril, 1851). COMMENT: Should Heterotropis Fitzinger, DIAGNOsIs: (1) Hemipenes bilobate with 1843, be demonstrated conclusively to be a distinctly divided sulci (also in polychrids); junior synonym of Stenocercus (and a senior (2) nasal concha fused to roof of nasal cham- ber. composed of a bony section and its confluent elongated CONTENT: Stenocercini, new tribe, and costal cartilage), of which at least the anterior pair is Tropidurini, new tribe. fused at the midline (Fritts, 1974; but see Etheridge, DISTIUBUTION: Most of tropical and sub- 1966). However, unlike "O." aculeatus and "O." cadu- tropical South America, excluding high ele- cus, in which the anteriormost inscriptional cartilages vations in the Andes and Patagonia. are well calcified and join seamlessly at the midline, at COMMENT: Further documentation and least "O." iridescens and "O." scapularis show anteri- characterization ofthe subfamilial taxonomy ormost inscriptional costal cartilages that variably close- of the Tropiduridae can be found in Frost ly approximate each other or are connected medially and Etheridge (1989). only by poorly chondrified connective tissue. In several "Stenocercus" (e.g., "S." apurimacus, "S." festae, "S." ornatus, "S." rhodomelas, and "S." trachycephalus), a STENOCERCINI, NEW TRIBE similar condition obtains of multiple, closely approxi- ?Heterotropides Fitzinger, 1843: 17. Type genus: mating postxiphisternal inscriptional ribs. For this rea- Heterotropis Fitzinger, 1843 (a nomen dubium) son Etheridge (1966) had considered species showing this (= Stenocercus Dumeril and Bibron, 1837?). condition to be members of a larger "Ophryoessoides" than that subsequently conceived of by Fritts (1974). DiAGNOsIs: (1) Superior fossa of quadrate However, only the more plesiomorphic inscriptional rib not enlarged (not penetrated by a quadrate pattern as seen in most other "Stenocercus" and in Proc- process of the squamosal); (2) alveolar shelf totretus is found in the type species of"Ophryoessoides," of mandible robust; (3) posterior process of "O." tricristatus (not examined by Etheridge, 1966, but the interclavicle anterior to contact with the who mentioned the possibility that this species might not be closely related to other "Ophryoessoides"). The sternum (4) hemipenial sheath muscu- long; reduced antehumeral and oblique neck folds of lature extensive (Arnold, 1984), lacking dor- "Ophryoessoides" are also found in Proctotretus (more sal accessory muscle; (5) gular scales imbri- so in P. azureus than P. pectinatus, which retains small cate posteriorly; (6) interparietal not enlarged antehumeral folds) and several species of "Stenocercus" or absent. (including those listed above). Proctotretus species also CONTENT: Stenocercus Dumeril and Bi- share certain apomorphic scale characteristics (e.g., dor- bron, 1837.5 solateral scale ridges) seen in some "Ophryoessoides." A synapomorphy of"Stenocercus" has yet to be suggested. Without digressing further into a revision of the Steno- 5 With the exception ofProctotretus Dumeril and Bi- cercini, it seems clear that "Ophryoessoides" and Proc- bron, 1837 (which has hyperossified phalanges and totretus are derived from "Stenocercus." Because several metacarpals [Proctotretus doellojuradoi not examined as workers describing new species in this tribe have ex- ofthis writing]), the other genera that compose the Steno- pressed some concern over generic definition in the group, cercini, "Ophryoessoides" Dumeril and Dumeril, 1851, and some have seemed inclined to publish revisions based and "Stenocercus" Dum6ril and Bibron, 1837, are not on comments in unpublished sections ofmy dissertation, natural taxa. "Stenocercus" (sensu Fritts, 1974) shares it seems best to me at this time to synonymize Ophryoes- the diagnosis ofthe Stenocercini and is paraphyletic with soides and Proctotretus with Stenocercus (stating Steno- respect to "Ophryoessoides" and Proctotretus (see be- cercus to have priority over Proctotretus under Article low). "Ophryoessoides" is characterized by features that 24 [Principle of the First Revisor] of the International are either of arguable polarity or homology, or variably Code ofZoological Nomenclature, 1985). Although many distributed in some "Stenocercus" and Proctotretus. These species pairs, or even monophyletic groups ofmore spe- include keeled ventral scales (also in Proctotretus azureus cies, could now be recognized generically, without a and so weak in "O." iridescens as to approach the con- cladogram of the entire tribe these actions would result dition in "S." trachycephalus) and more than two elon- only in the concomitant recognition ofunsupported and/ gate postxiphisternal inscriptional ribs (each ofwhich is or paraphyletic "taxa." 44 AMERICAN MUSEUM NOVITATES NO. 3033

80 60 40 Tropidurus Wied-Neuwied, 1825; Uranos- codon Kaup, 1826. 20 DISTRIBUTION: Tropical and subtropical South America, excluding northern and western Colombia and northeastern Venezuela, south to ca. 32°S (fig. 1). GENUS URANOSCODONKAUP, 1825 Uranoscodon Kaup, 1825: 590. Type species: La- certa superciliosa Linnaeus, 1758, by subsequent designation of Etheridge (1970a: 240). Ophryessa Boie, 1825: 1090. Type species: La- certa superciliosa Linnaeus, 1758, by subse- 20 quent designation of Fitzinger (1843: 16). Uraniscodon Boie, 1825: 1090. Unjustified emendation of Uranoscodon Kaup, 1825. Lophyrus Gray, 1827: 208 (not ofPoli, 1791 [Mol- lusca], or Oppel, 1811). Substitute name for Ur- anoscodon Kaup, 1825. Ophryoessa Wagler, 1830: 149. Unjustified emen- 40 dation of Ophryessa Boie, 1825. Ophyessa Gray, 1831: 39. Unjustified emendation of Ophryessa Boie, 1825.

20 DIAGNoSIS: (1) Skull highly elevated at lev- 100 80 60 40 el of orbits; (2) nutritive foramina of maxilla Fig. 37. Distribution of Stenocercini. not strikingly enlarged; (3) lingual process of dentary absent, not extending over lingual dentary process of coronoid; (4) angular not synonym of Ophryoessoides Dumeril and reduced; (5) medial centrale present; (6) Dumeril, 1851), the family-group name of "flash" marks on underside of thighs absent; Stenocercini would become Heterotropidini. (7) circumorbitals not distinct from other small supraorbital scales; (8) lateral fringe devel- TROPIDURINI BELL, 1843 oped on both sides of fourth toes; (9) hemi- Tropiduridae Bell, 1843: 1. Type genus: Tropidu- penes attenuate, without apical disks. rus Wied-Neuwied, 1825. CoNTENT: Uranoscodon superciliosus (Lin- Ptychosauri Fitzinger, 1843: 16. Type genus: Pty- naeus, 1758) (fig. 38). chosaurus Fitzinger, 1843 (= Plica Gray, 1831). DISTRIBUTION: Amazonian and Guianan Steirolepides Fitzinger, 1843: 17. Type genus: regions of South America (fig. 39). Steirolepis Fitzinger, 1843 (= Tropidurus Wied- ETYMOLOGY: Greek: ouranos (vault [= roof Neuwied, 1825). of the mouth]) + osco (mouth) + -odon DiAGNoSIS: (1) Superior fossa of quadrate (tooth): in reference to the presence of pter- enlarged (not penetrated by a quadrate pro- ygoid teeth. The gender is masculine. cess of the squamosal); (2) alveolar shelf of mandible somewhat eroded; (3) posterior pro- PLESIOMICROLOPHUS, NEW GENUS cess of the interclavicle anterior to contact with the sternum long; (4) elongate hemipenes TYPE SPECIES: Tropidurus koepekeorum (also in polychrids); (5) hemipenes with dor- Mertens, 1956. sal accessory muscle (also in polychrids); (5) DIAGNOSIS: (1) Skull not highly elevated at gular scales imbricate posterolaterally to lat- level oforbits; (2) nutritive foramina ofmax- erally (except in Tropidurus bogerti); (6) in- illa not strikingly enlarged; (3) lingual process terparietal enlarged (also in phrynosoma- of dentary present, extending over lingual tids). dentary process of coronoid; (4) angular not CoNTENT: Microlophus Dumeril and Bi- strongly reduced; (5) medial centrale present; bron, 1837; Plesiomicrolophus, new genus; (6) "flash" marks on underside of thighs ab- 1992 FROST: TROPIDURUS GROUP OF LIZARDS 45

I -,N

Fig. 38. Some members ofthe Tropidurini in life. Top: Uranoscodon superciliosus, KU 130218 (KU photo 3715). Photograph by M. L. Crump. Bottom: Plesiomicrolophus koepckeorum, J. R. Dixon photograph 105. sent; (7) circumorbitals distinct from other crolophus (see below), referencing the simi- small supraorbital scales, forming a single se- larity and possible phylogenetic propinquity ries; (8) lateral fringe not developed on both of this lineage to the Microlophus clade. The sides offourth toes; (9) hemipenes attenuate, gender is masculine. without apical disks. COMMENT: Although first described as a CONTENT: P. koepckeorum (Mertens, 1956) subspecies of Microlophus occipitalis, as (fig. 38). Tropidurus occipitalis koepckeorum, this spe- DISTRIBUTION: As for the single species: cies is almost completely plesiomorphic in foothills along the eastern side ofthe Sechura all features examined and could be consid- Desert, south to the Rio Shigiay, in Peru (fig. ered diagnostically "ancestral" to the clade 40). composed of Tropidurus and Microlophus. ETYMOLOGY: Greek: plesios (near) + Mi- My conjecture is that future work will show 46 AMERICAN MUSEUM NOVITATES NO. 3033

100 80 60 40 20 Fig. 39. Distribution ofMicrolophus and Ura- Fig. 40. Distribution ofPlesiomicrolophus and noscodon. Tropidurus. it to be the sister taxon of Microlophus, in "stem" species. That the diagnoses ofthe or- which case it could justifiably be considered ganisms constituting Plesiomicrolophus are a junior synonym of Microlophus. Although equivalent, or nearly so, to those in the an- Plesiomicrolophus has no unambiguous apo- cestral species is irrelevant, inasmuch as morphies, what evidence there is (Dixon and statements about ancestry apply to supraor- Wright, 1975) supports the notion that it is ganismal entities and not to organismal di- a single lineage. That the "genus" lacks apo- agnoses. morphies is strictly an artifact of the nomenclature convention. With other aspects ofcol- GENUS MICROLOPHUS or pattern, like narrow transverse dorsal bars, DUMERIL AND BIBRON, 1837 in this is gular spotting species likely apo- Microlophus Dumeril and Bibron, 1837: 334. Type morphic, although very similar (homolo- species: Microlophus lessonii Dumeril and Bi- gous?) spotting occurs in members ofthe Mi- bron, 1837 (= Stellio peruvianus Lesson, 1831), crolophus grayii complex (see below). by monotypy. Characterization ofthese patterns is extreme- Steirolepis Fitzinger, 1843: 72. Type species: Trop- ly difficult and has not been included in this idurus microlophus Wiegmann, 1835 (= Stellio analysis. peruvianus Lesson, 1831), by original designa- I agree with the principle discussed by Ax tion. (1985), that stem species do not survive lin- Craniopeltis Peters, 1871: 645. Type species: eage splitting, here exemplified by the fact Tropidurus bivittatus Peters, 1871, by mono- that the population that was ancestral to Ple- typy. Laemopristis Peters, 1871: 645. Type species: siomicrolophus, Microlophus, and Tropidu- Tropidurus occipitalis Peters, 1871, by mono- rus must be taken to currently be composed typy. of these three genera, not any one of the de- Aneuoporus Bocourt in Dumeril, Bocourt, and scendants, although it is possible that one of Mocquard, 1874: 215. Type species: Aneuopo- these may share a diagnosis with the ancestral rus occipitalis Bocourt, 1874, by monotypy. 1992 FROST: TROPIDURUS GROUP OF LIZARDS 47

A_fo/_-i/*- 1FA iXiirA

:t, X, , ' eJ,*J-'. .~ . ,

ll~>_v~ ~ tXJ~ f;.l ,,a

Fig. 41. Some members of the Tropidurini in life. Top: Microlophus albemarlensis, R. G. Zweifel photograph. Middle: M. occipitalis, KU 212667 (KU photo 8456), W. E. Duellman photograph. Bottom: M. theresioides, KU 162002 (KU photo 4940), W. E. Duellman photograph. 48 AMERICAN MUSEUM NOVITATES NO. 3033

~~~~~-a~~~~~~-

0 ..

-l ~- ,.,4,11~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

Fig. 42. Some members of the Tropidurini in life. Top: T. etheridgei, KU 160145 (KU photograph 4739), W. E. Duellman photograph. Bottom: Tropidurus melanopleurus, KU 182998 (KU photograph 6552 by D. C. Cannatella).

DiAGNosIs: (1) Skull not highly elevated at (Baur, 1890) (see comment below); M. grayii level oforbits; (2) nutritive foramina ofmax- (Bell, 1843) (see comment below); M. habelii illa not strikingly enlarged; (3) lingual process (Steindachner, 1876); M. heterolepis (Wieg- of dentary present, extending over lingual mann, 1834); M. occipitalis (Peters, 1871) (fig. dentary process of coronoid; (4) angular not 41); M. pacificus (Steindachner, 1876) (see strongly reduced; (5) medial centrale present; comment below); M. peruvianus Lesson, (6) "flash" marks on underside of thighs ab- 1831; M. quadrivittatus (Tschudi, 1845); M. sent; (7) circumorbitals distinct from other stolzmanni (Steindachner, 1891); M. tara- small supraorbital scales, forming a single se- pacensis (Donoso-Barros, 1966) (not seen but ries; (8) lateral fringe not developed on both provisionally allocated here); M. theresiae sides offourth toes; (9) hemipenes with apical (Steindachner, 1901); M. theresioides (Dono- disks. so-Barros, 1966) (fig. 41); M. thoracicus CONTENT: Microlophus albemarlensis (Tschudi, 1845); M. tigris (Tschudi, 1845); (Baur, 1890) (fig. 41) (see comment below); M. yanezi (Ortiz-Zapata, 1980). M. atacamensis (Donoso-Barros, 1966); M. DISTRIBUTION: Galapagos Islands; South bivittatus (Peters, 1871); M. delanonis (Baur, America west of the Andes from southern 1890) (see comment below); M. duncanensis Ecuador to northern Chile; east of the con- 1992 FROST: TROPIDURUS GROUP OF LIZARDS 49

Fig. 43. Tropidurusflaviceps, R. W. McDiarmid photograph 18872. tinental divide only in the Huancabamba De- islands from the mainland, and M. grayii rep- pression Region of northern Peru (fig. 39). resenting the other. However, ifthe assump- ETYMOLOGY: Greek: mikros (small) + lo- tion of clocklike molecular evolution is not phos (crest); in reference to the reduced dorsal made for purposes of data analysis, the cla- crest in the type species, M. peruvianus. The distic structure ofthe allozymic data is murky. gender is masculine. An alternative would be to recognize all diag- COMMENT: As in most archipelago species nosable allopatric populations on the islands complexes, the current taxonomy of Gala- as species (sensu Frost and Hillis, 1990). The pagos Microlophus has inherent difficulties. inter- and intraisland variation in squama- Each island has its own population(s) that tion and coloration documented by Van Den- differs in some respect from all other island burgh and Slevin (1913) makes this alterna- populations (Wright, 1983). That some ofthe tive attractive. However, although I think that recognized species (e.g., Microlophus albe- a taxonomic treatment of this complex is marlensis) are found on several islands and needed, because this group was not the focus are lumped together under one binomial be- of this study I resist taking the obvious step cause ofoverall similarity (Van Denburgh and ofrecommending a three-species or multiple- Slevin, 1913) may reflect grouping by plesio- species model of Galapagos lava lizard tax- morphy rather than on an understanding of onomy without additional study. the historical relationships of these island forms. Work by Wright (1983) on allozyme GENUS TROPIDURUS distances, although not dealing explicitly with WIED-NEUWIED, 1825 species distinction, has made it arguable that Tropidurus Wied-Neuwied, 1825: 131. Type spe- only three species, composing two mono- cies: Stellio torquatus Wied-Neuwied, 1820, by phyletic groups, could be recognized: Micro- subsequent designation of Fitzinger (1843: 17). lophus habelii and M. bivittatus possibly rep- Uracentron Kaup, 1826: 88. Type species: Ura- resenting the fruits of one invasion of the centron brevicaudatum Kaup, 1826 (= Lacerta 50 AMERICAN MUSEUM NOVITATES NO. 3033

Fig. 44. Top: Tropidurus plica, L. J. Vitt photograph. Bottom: T. umbra, R. W. McDiarmid photograph 13411.

azurea Linnaeus, 1758), by subsequent desig- Strobilurus torquatus Wiegmann, 1834b, by nation of Fitzinger (1843: 17). monotypy. Doryphorus Cuvier, 1829: 34. Type species: Stellio Hypselophus Wiegmann, 1835: 289. Substitute brevicaudatus Latreille, 1802 (= Lacerta azurea name for Hypsibatus Wagler, 1830. Linnaeus, 1758), by monotypy. Hypselopus Gravenhorst, 1837: 717. Unjustified Hypsibatus Wagler, 1830: 150. Type species: La- emendation of Hypselophus Wiegmann, 1835. certa umbra Linnaeus, 1758, by subsequent des- Uperanodon Dumeril and Bibron, 1837: 247. Type ignation of Fitzinger, 1843: 16. species: Lophyrus ochrocollaris Spix, 1825 (= Platynotus Wagler, 1830: 146. Type species: Aga- Lacerta umbra Linnaeus," 1758), by monotypy. ma semitaeniata Spix, 1825, by monotypy. Pre- Ptychosaurus Fitzinger, 1843: 59. Type species: occupied by Platynotus Fabricius, 1801 (Cole- Hypsibatus punctatus Dumeril and Bibron, 1837 optera). (= Lacerta plica Linnaeus, 1758), by original Plica Gray, 1831: 40. Type species: Lacerta plica designation. Linnaeus, 1758, by subsequent designation of Ptychopleura Fitzinger, 1843: 59. Type species: Etheridge (1970a: 241). Hypsibatus plica Wagler, 1830 (= Lacerta plica Strobilurus Wiegmann, 1834b: 18. Type species: Linnaeus, 1758), by original designation. 1992 FROST: TROPIDURUS GROUP OF LIZARDS 51

Taraguira Gray, 1845: 219. Type species: none ourus (tail), referring to the keeled squama- designated. tion on the tails of most species. Hyperanodon Agassiz, 1847: 190. Substitute name for Uperanodon Dumeril and Bibron, 1837. Tapinurus Amaral, 1933: 65. Type species: Tapi- REFERENCES nurus scutipunctatus Amaral, 1932 (= Agama Adams, E. N. semitaeniata Spix, 1825), by original designa- 1972. Consensus techniques and the compar- tion. ison of taxonomic trees. Syst. Zool. 21: 390-397. DIAGNOSIS: (1) Skull not highly elevated at Agassiz, A. level of orbits (except in T. umbra); (2) nu- 1847. Nomenclator zoologici universalis.... tritive foramina of maxilla strikingly en- Soloduri, Jent and Gassman. viii, 393 larged; (3) lingual process of dentary present, pp- extending over lingual dentary process of cor- Amaral, A. onoid; (4) angular strongly reduced; (5) me- 1933 "1932." Estudos sobre lacertilios neotro- dial centrale absent; (6) "flash" marks on un- picos. I. Novos generos e especies de derside of thighs present, yellow to black lagartos do Brasil. Mem. Inst. Butantan (obscured or lost in T. azureus and T. 7(1932): 51-74. flavi- Andersson, L. G. ceps); (7) circumorbitals distinct from other 1918. New lizards from South America. Col- small supraorbital scales, in one or two rows; lected by Nils Holmgren and A. Roman. (8) lateral fringe not developed on both sides Ark. Zool. 11(16): 1-9. offourth toes; (9) hemipenes attenuate, with- Arnold, E. N. out apical disks. 1984. Variation in the cloacal and hemipenial CoNTENr: T. amathites Rodrigues, 1984 muscles of lizards and its bearing on (not seen but provisionally allocated here); T. their relationships. Symp. Zool. Soc. azureus (Linnaenus, 1758); T. bogerti Roze, London 53: 47-85. 1958; T. cocorobensis Rodrigues, 1987; T. Ax, P. divaricatus Rodrigues, 1986 seen but 1985. Stem species and the stem lineage con- (not cept. Cladistics 1: 279-287. provisionally allocated here); T. erythroceph- Baur, G. alus Rodrigues, 1987; T. etheridgei Cei, 1982 1890. Das Variieren der Eidechsen-Gattung (fig. 42); T. flaviceps (Guichenot, 1855) (fig. Tropidurus auf den Galapagos Inseln 43); T. helenae (Manzini and Abe, 1990) (not und Bemerkungen uiber den Ursprung seen but clearly allocated here); T. hispidus der Inselgruppe. Biol. Centralblatt. 10: (Spix, 1825); T. hygomi Reinhardt and Liut- 475-483. ken, 1861; T. insulanus Rodrigues, 1987; T. 1892. Das Variieren de Eidechsen-Gattung itambere Rodrigues, 1987; T. lumarius (Don- Tropidurus auf den Galapagos-Inseln. nelly and Myers, 1991); T. melanopleurus In Festschrift zum Siebenzigsten Ge- Boulenger, 1902 (fig. 42); T. montanus Ro- burtstage RudolfLeukarts, pp. 259-277. Leipzig. drigues, 1987; T. mucujensis Rodrigues, 1987; Bell, T. T. nanuzae Rodrigues, 1981; T. oreadicus 1843. Zoology of the voyage of the H.M.S. Rodrigues, 1987; T. pinima (Rodrigues, Beagle, under the command of Captain 1984); T. plica (Linnaeus, 1748) (fig. 44); T. Fitzroy, R. N., during the years 1832 to psammonastes Rodrigues et al., 1988 (not 1836. Edited and superintended by seen but provisionally allocated here); T. Charles Darwin ... naturalist to the ex- semitaeniatus (Spix, 1825); T. spinulosus pedition. Part 5. Reptiles. London: (Cope, 1862); T. strobilurus, new name (for Smith, Elder & Co, vi + 51 pp., atlas. Strobilurus torquatus Wiegmann, 1834; the Berthold, A. A. combination Tropidurus torquatus is preoc- 1859. Einige neue Reptilien des Akad. zoolog. cupied; see next Museums in Gottingen. Nachr. Georg- entry); T. torquatus Wied- Augustus Univ. K. Ges. Wiss. Gottin- Neuwied, 1825; T. umbra (Linnaeus, 1758) gen 1859(17): 179-181. (fig. 44). Bocourt, M. F. DISTRIBUTION: Tropical to temperate South 1874. In A. Dumeril, M. F. Bocourt, and F. America east of the Andes (fig. 40). Mocquard. 1870-1909. Mission scien- ETYMOLoGY: Greek: tropido- (keeled) + tifique au Mexique et dans l'Amerique 52 AMERICAN MUSEUM NOVITATES NO. 3033

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1891. Ueber neue und seltene Lacertiden aus Wagler, J. G. den herpetologischen Sammlung des k. 1830. Natiurliches System der Amphibien, mit k. naturhistorisches Hofsmuseums. Ann. vorangehender Classification der Sau- k. k. Hofmus. Wien 6: 371-378. gethiere und Vogel. Miinchen, Stuttgart, 1901. Herpetologische und ichthyologische and Tubingen: J. G. Cotta, vi + 354 pp. Ergebnisse einer Reise nach Siidameri- Watrous, L. E., and Q. D. Wheeler ka mit einer Einleitung von Therese 1981. The outgroup comparison method of Prinzessin von Baiern. Anz. Akad. Wiss. character analysis. Syst. Zool. 30: 1-11. Wien 38: 194-196. Werner, F. 1902. Herpetologische und ichthyologische 1899. Beschreibung neuer Reptilien und Ba- Ergebnisse einer Reise nach Siidameri- trachier. Zool. Anz. 22: 479-484. ka, mit einer Einleitung von Therese 1900. Reptilien und Batrachier aus Peru und Prinzessin von Bayern. Denkschr. Akad. Bolivien. Abh. Ber. K. Zool. Anthropol. Wiss. Wien 72: 89-148. Ethnol. Mus. Dresden 9: 1-4. Stejneger, L. Wied-Neuwied, M., Prinz zu 1901. An annotated list of batrachians and 1820-21. Reise nach Brasilien en den Jahren reptiles collected in the vicinity ofGuai- 1815 bis 1817. 2 vols. Frankfurt a. M.: ra, Venezuela, with description of two H. L. Br6nner. new species of snakes. Proc. U.S. Natl. 1825. Beitriige zur Naturgeschichte von Bra- Mus. 24: 179-192. silien, vol. 1. Weimar: Gr. H. S. priv. Stevens, P. F. Landes-Industrie-Comptoirs. 1980. Evolutionary polarity ofcharacter states. Wiegmann, A. F. A. Annu. Rev. Ecol. Syst. 11: 333-358. 1834a. Beitrage zur Zoologie gesammelt aufei- Swofford, D. L. ner Reise um die Erde, von Dr. F. J. F. 1989. PAUP-Phylogenetic Analysis Using Meyen. Siebente Abhandlung. Amphi- Parsimony. Version 3.01. User's Man- bien. Nova Acta Acad. Caesar. Leop. ual. Privately Published. Carol., Halle 17: 185-268. 1834b. Herpetologica Mexicana seu descriptio Tschudi, J. J. von amphibiorum Novae Hispaniae. Pars 1845. Reptilium conspectum quae in republi- prima. Saurorum species. Berlin: Lii- ca Peruana reperiuntur et pleraque ob- deritz, vi + 54 pp. servata vel collecta sunt in itinere. Arch. 1835. Bericht iuber die Fortschritte der Zool- Naturgesch. 11: 150-170. ogie in Jahre 1834. Amphibien. Arch. Valdivieso, D., and J. R. Tamsitt Naturgesch. 1: 273-296. 1963. Records and observations on Colom- Wiley, E. 0. bian reptiles. Herpetologica 19: 28-29. 1979. An annotated Linnaean hierarchy, with Van Denburgh, J., and J. R. Slevin comments on natural taxa and compet- 1913. Expedition of the California Academy ing systems. Syst. Zool. 28: 308-337. of Sciences to the Galapagos Islands, 1981. Convex groups and consistent classifi- 1905-1906. IX. The Galapagoan lizards cations. Syst. Bot. 6: 346-358. of the genus Tropidurus with notes on Wright, J. W. iguanas of the genera Conolophus and 1983. The evolution and biogeography of the Amblyrhynchus. Proc. California Acad. lizards of the Galapagos Archipelago: Sci., ser. 4, 2: 132-202. evolutionary genetics of Phyllodactylus Vanzolini, P. E., and N. Gomes and Tropidurus populations. In R. I. 1979. On Tropidurus hygomi: redescription, Bowman et al. (eds.), Patterns of evo- ecological notes, distribution and his- lution in Galapagos organisms, pp. 123- tory (Sauria, Iguanidae). Pap. Avulsos 155. AAAS Symposium vol., San Fran- Zool., Sao Paulo 32: 243-259. cisco. 58 AMERICAN MUSEUM NOVITATES NO. 3033

APPENDIX 1 SPECIMENS EXAMINED

Abbreviations for names of collections are: AMNH American Museum of Natural History; BMNH The Natural History Museum, London; CAS California Academy of Sciences; KU Museum of Natural History, University of Kansas; LACM Natural History Museum of Los Angeles County; LSUMZ Mu- seum ofNatural Science, Louisiana State University; MAN Mark A. Norell private osteology collection; MCZ Museum of Comparative Zoology, Harvard University; MPEG Museu Paraense Emilio Goeldi, Belem, Brazil; MVZ Museum ofVertebrate Zoology, University ofCalifornia; REE Richard E. Etheridge osteology collection; RWM Roy W. McDiarmid field series (to be accessioned into USNM); SDSNH San Diego Natural History Museum; TCWC Texas Cooperative Wildlife Collection, Texas A&M Uni- versity; UMMZ Museum ofZoology, University of Michigan; UNM Museum of Southwestern Biology, University of New Mexico; USNM National Museum of Natural History, Smithsonian Institution. Abbreviations for specimens are: A Alizarin stained skeleton, AA Alcian blue-alizarin red stained postcranial skeleton, CAA complete alcian blue-alizarin red skeleton, D dry skeleton, S dry skull, SN skin, H hemipenial preparation or dissection, AL alcoholic specimen, X X-ray photograph. All special preparations (e.g., injected hemipenes, skeletons) are listed but AMNH and KU alcoholic material is not noted. Additionally, although much iguanian material of virtually all genera (except for chamae- leonids) has been examined at one time or another, only material representing my ingroup and immediate outgroups is here listed. Tropidurini M. theresioides: KU 162012 (AA, S), 162018 (AA, S); LACM 134136-41 (AL). M. thoracicus: KU Microlophus albemarlensis: CAS 11153 (S); 163721 (AA, S), 163724 (AA, S); LACM 48914 LACM 106243-49 (AL), 106250 (AA, S), 106251- (AL), 48917 (AL), 48922 (AL). M. tigris: KU 60 (AL), 106261 (AA, S), 196261 (D); MCZ 28475 163752 (H), 163753 (AA, S), 163756 (AA, S); (D), 36928 (D); MVZ 77323 (D), 77467 (D), 77469 SDSNH 47031 (D). M. yanezi: MVZ 92938-39 (D); SDSNH 7211-12 (D), 7220 (D). M. ataca- (D, SN), 92945 (D, SN). Plesiomicrolophus mensis: KU 161983 (AA, S, H), 161986 (AA, S). koepckeorum: LACM 49080-81 (AL), 49082 M. bivittatus: AMNH 20499 (H); LACM 106302 (CAA), 49083 (AL, H), 49084 (AL), 49085 (AA, (AL, H), 106303 (AA, S), 106305-06 (AL), 106307 S), 49086-87 (AL), 49088 (AA, S), 49089-104 (AA, S), 106308 (AL); SDSNH 7192 (2) (D). M. (AL), 109569 (AL), 122584 (AL), 122588 (AL), delanonis: AMNH 92731 (AL, H), MCZ 9204 (D), 122590-91 (AL); TCWC 28687-88 (AL, H), 28697 36909 (2) (D), 36910 (D); SDSNH 10065 (D), (AL, H), 28700 (AL, H). Tropidurus azureus: 10073 (D), 10241 (D), 22108 (D). M. duncanensis: AMNH 60330 (S, AA), 61014 (S, AA); MAN 47 CAS 12202 (D); SDSNH 10194 (D). M. grayii: (D); UMMZ 129418 (A). T. bogerti: RWM 11659- CAS 11620 (D). M. habelii: LACM 106380-87 61 (AL), 11662 (D, SN), 11663 (CAA), 11664 (AL, (AL), 106388 (AL, H), 106389-90 (AL), 106391 H). T. cocorobensis: MCZ 172946-47 (AL, X). T. (AA, S), 106392-93 (AL), 106394 (AA, S), 106395- erythrocephalus: MCZ 172948 (S, AA), 172949- 96 (AL). M. heterolepis: AMNH 65331-42 (AL, 50,172952,172954 (AL, X). T. etheridgei: AMNH X). M. occipitalis: KU 142714 (AA, S, SN), 142721 172948 (S, AA), 17949-50 (AL, X); KU 160145 (AA, S), 142735 (CAA), 163630 (H); LACM 48870 (H), 186102 (AA, S), 186113 (AA, S); LACM 73991 (AL), 48874 (AL, H), 48902 (AL), 48904-05 (AL); (AL, H); UNM 98982 (D). T. flaviceps: AMNH REE 668 (D), 1847 (D), 1860 (D). M. pacificus: 71101-02 (AL, X); KU 126781 (H), 175317 (AA, CAS 12480 (D); MCZ 28495 (2) (D); SDSNH S). T. hispidus: CAS 49549 (D); KU 167508 (AA, 10224 (D), 10230 (D). M. peruvianus: KU 134673 S, H), 167513 (AA, S); MCZ 3133 (D), 3438 (D), (AA, 2), 134674 (AA, S, H), 134679 (H), 134695 38536 (D), 43869 (D), 49529 (D); SDSNH 34882- (AA, S), 163640 (H), 164055-56 (D); LACM 49016 83 (D); USNM 159200-05 (AL, X). T. hygomi: (AL, H), 49018 (AL, H), 49021 (AL), 49051 (AL, AMNH 37542 (S, AA), 37538-41 (AL, X); USNM H), 49053 (AL, H); SDSNH 30843 (D), 30893 (D), 209643 (AL, X), 209644 (AA, S), 209645-46 (AL, 30911 (D). M. quadrivittatus: REE unnumbered X). T. insulanus: MCZ 172879-82 (AL, X). T. (3) (AL, X). M. stolzmanni: KU 134701 (H), itambere. USNM 148773 (S, AA), 148775-77 (AL, 134704 (AA, S), 134731 (CAA), 134747 (AA, S); X); MCZ 172883 (AL, X), 172885-87,172892 (S, LACM 49105 (AL, H), 49119 (AL, H), 49121 (AL, AA). T. lumarius: AMNH 136177 (S, AA), 136176 H), 49129 (AL, H), 49132 (AL, H). M. theresiae: (H). T. melanopleurus: KU 136367 (D), 136370- LACM 49060-61 (AL), 49064 (AL), 49070 (AL), 71 (AA, S), 136374 (D), 183469 (H); REE 2612 49076-77 (AL), 122696 (AL), 122702 (AL), (D), 2616 (D). T. montanus: USNM 218204-9 122706 (AL), 122707-08 (AA, S), 122710 (AL). (AL, X), 218210 (S, AA); MCZ 172893-94, 1992 FROST: TROPIDURUS GROUP OF LIZARDS 59

172896,172901-02. T. mucujensis: MCZ 172944- 163602 (AA, S). S. cupreus: KU 133974 (AA, S), 45 (AL, X). T. nanuzae: MCZ 160243 (AA, S, 133976 (AA, S); MCZ 43789 (D). S. empetrus: SN), 160244-45 (AL); USNM 213514-15 (AL, KU 134401 (AA, S), 134403 (AA, S), 134421 H). T. oreadicus: UMMZ 56805 (S, AA), 188632- (CAA), 181909 (H). S. erythrogaster: MCZ 36877 33 (S); USNM 188632-33 (D). T. pinima: AMNH (S); UMMZ 149107 (D). S. festae: KU 134588 131862-63 (AL, X). T. plica: AMNH 61314 (D), (H), 134595 (AA, S), 134603 (AA, S), 141161 (H). 85313 (D), 107590 (AL); KU 117088 (H), 167499 S. formosus: KU 134110 (AA, S), 134112 (H); (AA, S); MAN 76 (D); MCZ 6100 (D), 9001 (D), MCZ 11295 (S). S. guentheri: KU 147319 (AA, 43865 (D); UMMZ 149129 (D). T. semitaeniatus: S), 147326 (AA, S), 147347 (CAA), 179426 (H), AMNH 131864-66 (AL, X); CAS 49386-88 (AL), 202940 (H); MCZ 8418 (D), 8423 (D), 8427 (D). 49455 (AL), 49468-69 (AL), 49471-72 (AL), S. humeralis: KU 121136 (H), 134001 (AA, S), 49473 (D), 49474-76 (AL); LSUMZ 39519 (AA, 134004 (AA, S). S. iridescens: AMNH 21993 (AA, S); MCZ 79805 (D), 131890-91 (D). T. spinulosus: S), 112989 (AA, S), 112990 (AA, S); MCZ 8412 AMNH 101490-91 (S, AA); CAS 49843 (D); KU (S), 84162 (D), 84165 (D), 84167 (D). S. mar- 97853 (H), 97856 (AA, S); LACM 126315-16 (AL), moratus: UMMZ 149276 (D). S. melanopygus: 126318 (AL), 126321 (AA, S); UNM-ALA 237 KU 134058 (AA, S), 134075 (AA, S). S. nigro- (D); UNM 99121 (AL, S), 99128 (AL); USNM maculatus: KU 134089 (AA, S), 134092 (AA, S), 125166 (AL, S), 126016 (AL), 126029 (AL, S), 134105 (H); MCZ 18767 (D). S. ochoai: KU 126703 (D), 126705 (D), 126707 (D), 126941 (D), 133876 (CAA), 133878 (AA, S), 133884 (AA, S, 128312 (AL, S), 128987 (D). T. strobilurus: BMNH H), 139267 (H). S. orientalis: KU 134452 (AA, 1903.10.16.23 (D); MCZ 154211 (AL, X), 133243 S), 134460 (AA, S), 134464 (CAA). S. ornatissi- (D); MPEG unnumbered (AL, H, X). T. torquatus: mus: KU 134351 (AA, S), 134360 (AA, S). S. AMNH 62148 (S, AA); KU 128205 (H); MVZ ornatus: KU 121128 (AA, S), 134128 (AA, S), 92970 (D, SN); USNM 98593-98 (AL), 207683- 134130 (H), 134155 (CAA). S. praeornatus: KU 85 (AL, X). T. umbra: AMNH 61239 (D), 61436 134225 (H), 134229 (AA, S). S. rhodomelas: KU (D); KU 135267 (D), 146659 (AA, S, H), 147946 142699 (H), 152184 (AA, S), 152186 (AA, S). S. (H); LACM 49140 (AL, H); MCZ 152184 (D). roseiventris: KU 134156 (H), 172196 (AA, S). S. Uranoscodon superciliosus: AMNH 61304 (D); KU scapularis: AMNH 56770 (AA, S), 56777 (AA, S). 128215 (D), 128216 (H), 128218 (H), 135269 (D); S. simonsii: KU 134163 (H). S. trachycephalus: LACM 44474 (AL, H); MCZ 9318 (D), 58340 (D); AMNH 131223 (S, AA), 131227 (S, AA); MCZ REE 16600; UMMZ 149312 (D); USNM 202682 17144 (D). S. variabilis: KU 134178 (H), 134198 (S). (AA, S), 134213 (AA, S). S. varius: KU 121135 (H), 134563 (H), 142704 (AA, S). Stenocercini Leiocephalinae Stenocercusaculeatus: KU 121093 (AA, S); MCZ 8061 (S); UMMZ 149102 (D). S. apurimacus: KU Leiocephalus carinatus: UMMZ 149103-04 (D); 134244 (H), 134284 (AA, S), 134288 (H), 134306 USNM 81709 (D). L. greenwayi: UMMZ 149108 (CAA). S. boettgeri: KU 134011 (H), 134014 (AA, (D). L. inaguae: UMMZ 149133 (D). L. loxo- S); MCZ 45843 (D). S. caducus: AMNH 37907 grammus: KU 192293 (D); UMMZ 149134 (D). (CAA); MCZ 20625-26 (AL); UMMZ 149105-06 L. psammodromus: UMMZ 149109 (D). L. punc- (D). S. chrysopygus: KU 133895 (AA, S), 133906 tatus: UMMZ 149110 (D). L. raviceps: UMMZ (AA, S), 134315 (H); MCZ 45832. S. crassicau- 149111 (D). L. schreibersi: KU 93358 (CAA). L. datus: AMNH 23132 (D); KU 133959 (AA, S), vittatus: CAS 39304 (S). 60 AMERICAN MUSEUM NOVITATES NO. 3033

APPENDIX 2 DATA MATRIX

1 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 7777777 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567 Ancestor 0000000000 0000?00000 ?000000000 0000000000 0000?00??0 0000?00000 0000000000 0000000 Uranoscodon 0101100000 1100000010 0110001101 0000001000 00011114?0 1000?01010 0101001002 3000010 Tropidurus atacamensis 0000000000 1110000000 0000001100 0010000000 00110001?O 0010?00001 0100001000 1111000 T. peruvianus 0000000000 1110000000 0000001100 0010000000 00110001?O 0010?00001 0100001000 1011000 T. koepckeorum 0000000000 1110000000 0000001100 0010000000 00010000?O 0000?00000 0100001000 0000000 T. occipitalis 0000000000 1110000000 0000001100 0010000000 00110000?0 0000?00000 0100001000 0000000 T. stolzmanni 0000000000 1110000000 0000001100 0010000000 00110010?0 0000?00000 0100001000 0000000 T. bivittatus 0000000000 1110001000 0000001100 0010000000 00110000?0 0000?00000 0100001000 0000000 T. nanuzae 1000001000 1211112001 0100011101 0021100000 0101000211 0000?00000 0100001000 0000000 T. cocorobensis 1000001000 12??????01 0000101101 0021100000 02?1?00200 0000?00000 0100001000 0100000 T. hygomi 1000001000 1211112001 0000101101 0021100000 0201100200 0000?00000 0100001000 0100000 T. erythrocephalus 1000001000 1211112001 0000101101 0021100000 02???00210 0001010000 0100001000 0100000 T. hispidus 1000001000 1211112001 1000101101 1021100000 0201100210 0001100000 0100001000 0100000 T. insulanus 1000001000 12??????01 0000101101 1021100000 0201?00210 0001000000 0100001000 1100000 T. itambere 1000001000 1211112001 0000101101 0021100000 0201100210 000??10000 0100001100 0100000 T. montanus 1000001000 1211112001 0000101101 1021100000 0201100220 0001110000 0100001000 0100000 T. mucujensis 1000001?00 12??????01 0000101101 1021100000 02???00210 1001110000 0100001000 0100000 T. oreadicus 1000001000 1211112001 0000101101 1021100000 0201100220 0001000000 0100001000 0100000 T. torquatus 1000001000 1211112001 0000101101 1021100000 0201100200 1001110000 0100001000 0100000 T. bogerti 1000011000 1211012001 0001101011 1021200010 12011001?0 1101111000 00?0001000 2100000 T. melanopleurus 1000001000 1211112001 0000101111 1021200000 11011013?0 1100?00010 0110101000 2010000 T. spinulosus 1000001000 1211110001 0000101011 1021200000 01011023?0 1100?00110 0110101000 2000000 Tapinurus semitaeniatus 1010021000 1211112111 1001101121 0121200010 12011001?0 1101110000 0110001000 2110000 Plica plica 1000101101 1211112011 1110101101 1021200000 21011123?2 1100?01110 0121101011 2000000 P. umbra 1101101101 1211112001 1111001101 1021200000 20011023?0 0000?01110 0111101012 3000010 Strobilurus 1000101100 1211112011 1011101101 1021200100 21011000?0 1100?00010 0110101001 2000101 Uracentron azureum 1000101110 1211110111 1011001000 1021210101 20011111?0 0100?01000 1010111101 3110200 U. flaviceps 1000101100 1211110111 1110001100 1021210101 20011111?0 1100?01000 1010111101 3100200 1 992 FROST: TROPIDURUS GROUP OF LIZARDS 61

APPENDIX 3 Changes by Transformation Series for Figure 33 Character changes noted by a double dagger (t) are of equally parsimonious placement elsewhere. In the case of terminal taxa even though the character assignment is not equivocal, the plesiomorphic member of the transformation pair may have alternative placement in the topology of the tree. Trans- formation Series denoted with parentheses ( ) are unpolarized (additive) in the analysis; those surrounded by brackets [ ] are unordered (nonadditive). Trans. Character series ci Steps change Along stem 1 1.000 0 - 1 5 2 0.500 0-. 1 Uranoscodon superciliosus Plica umbra 3 1.000 0 1 Tapinurus semitaeniatus 4 0.500 0-. 1 Uranoscodon superciliosus 0-. 1 Plica umbra 5 0.500 0 - 1 Uranoscodon superciliosus 0-. 1 16 6 1.000 14 1 - 2 Tapinurus semitaeniatus 7 1.000 0 - 1 5 8 1.000 0 - 1 16 9 1.000 0 - 1 Uracentron azureum 10 1.000 0 1 17 11 1.000 1 0. 1 1 12 1.000 1 0 - 1 1 1 1 2 5 13 1.000 1 0 1 2 14 1.000 0 - 1 5 15 0.500 0 - 1 5 1 1 0 T. bogerti 16 1.000 1 0-. 1 5 [17] 0.500 0-. 1 T. bivittatus 0 - 2 5 1 2 - 0 T. spinulosus 1 2 - 0 19 18 0.500 1 0 - 1 Tapinurus semitaeniatus 0-. 1 19 19 0.250 0-. 1 Uranoscodon superciliosus Tapinurus semitaeniatus 0 1I 16 1 - 0 Plica umbra 20 1.000 1 5 1 (21) 0.333 0-. T. 1 hispidus 0-. Tapinurus 1 semitaeniatus 0-. 16 1 22 0.250 0-. 1 Uranoscodon superciliosus 0-. T. 1 nanuzae 0-. 17 0-. 1 1 Uracentron flaviceps 23 0.500 0-. 1 Uranoscodon superciliosus 0-. 16 24 0.250 0- 1 14 16 1 - 0 Plica plica 1 - 0 Uracentron flaviceps 62 AMERICAN MUSEUM NOVITATES NO. 3033

APPENDIX 3 Changes by Transformation Series for Figure 33-(Continued) Trans. Character series ci Steps change Aloniig stem 25 0.333 1 0 - 1 6 1 1 - 0 Plica umbra 1 1 - O 19 26 1.000 1 0 - 1 T. nanuzae 27 1.000 1 0 - 1 1 28 0.250 1 0 1 1 1 1 0 T. bogerti 1 1 -b 0 T. spinulosus 1 1 10 Uracentron azzureum 29 0.667 1 0 1 13 1 1 - 2 Tapinurus sennitaeniatus 1 1 -0 16 30 0.333 1 0 - 1 Uranoscodon superciliosus 1 0 1 5 1 1 -0 19 31 0.500 1 0 1 9 1 1 - 0 Tapinurus sennitaeniatus 32 1.000 1 0 - 1 Tapinurus sennitaeniatus 33 1.000 1 0 1 2 1 1 2 5 34 1.000 1 0 1 5 35 1.000 1 0 1 5 1 1 2 13 36 1.000 1 0 1 19 37 1.000 1 0 1 Uranoscodon superciliosus 38 1.000 1 0 1 18 39 1.000 1 0 1 14 40 1.000 1 0 1 19 [41] 0.667 1 0 ijt 13 1 1 0 T. spinulosus 1 1 -2 16 [42] 0.400 1 0 1it 5 1 1 2t 6 1 2 1 15 1 1 - 0 Plica umbra 1 10 19 43 1.000 1 0 - 1 3 44 1.000 1 0 - 1 1 (45) 0.500 1 0 - 1 Uranoscodon 6superciliosus 1 0 - 1 6 46 0.333 1 0 - 1 Uranoscodon superciliosus 1 0 1 Plica plica 1 0 - 1 19 47 0.333 1 0 - 1 Uranoscodon superciliosus 1 0 -: 1 T. stolzmanni 1 0 - 1 15 1 1 - 2 T. spinulosus 1 1 - 2 17 1 1 -- 0 Strobilurus torrquatus 1992 FROST: TROPIDURUS GROUP OF LIZARDS 63

APPENDIX 3 Changes by Transformation Series for Figure 33-(Continued) Trans. Character series ci Steps change Along stem [48] 0.571 1 0 - 4t Uranoscodon superciliosus 1 0 - 1 4 1 0- 2 5 1 2- 1t 13 1 1- 3t 15 1 3- Ot 18 1 0 - itt 19 [49] 0.500 1 0 - 1 7 1 1 - 2 T. montanus 1 1 0 T. torquatus 1 1 -a 2 T. oreadicus [50] 1.000 1 0 - 1 T. nanuzae 1 0 - 2 Plica plica 51 0.250 1 0 - 1 Uranoscodon superciliosus 1 0 - 1 12 1 1 - 0 Plica umbra 1 1 0 Uracentron azureum 52 0.500 1 0 1 13 1 1 - 0 Plica umbra 53 1.000 1 0 1 4 54 0.500 1 0- 1 7 1 1 -0 15 (55) 1.000 1 0 - 1 10 56 0.333 1 0- 1 8 1 0 1 11 1 1-0O 15 57 0.250 1 0 1 Uranoscodon superciliosus 1 0 1 T. bogerti 1 0-1*it 16 1 1 - 0 Strobilurus troquatus 58 0.500 1 0 1 T. spinulosus 1 0 - 1 17 59 0.333 1 0 1 Uranoscodon superciliosus 1 0 - 1 15 1 1- 0 19 60 1.000 1 0 - 1 4 61 1.000 1 0 - 1 19 62 0.333 1 0-1 1 1 1 -b 0 T. bogerti 1 1-0O 19 63 1.000 1 0 1 12 1 1 - 2 Plica plica 64 0.500 1 0 - 1 Uranoscodon superciliosus 1 0 - 1 17 65 1.000 1 0- 1 15 66 1.000 1 0 - 1 19 67 1.000 1 0 - 1 1 68 0.500 1 0 - 1 T. itambere 1 0 - 1 19 69 1.000 1 0-1 17 64 AMERICAN MUSEUM NOVITATES NO. 3033

APPENDIX 3 Changes by Transformation Series for Figure 33-(Continued) Trans. Character series ci Steps change Along stem 70 0.500 2 0 - 2 Uranoscodon superciliosus 1 0 1 16 1 1 - 2 Plica umbra [711 0.500 1 0 - 3 Uranoscodon superciliosus 1 0 1 4 1 0 - 2 13 1 2 -+ 3 Plica umbra 1 2 - 3 19 1 0 - 1 T. insulanus 72 0.250 1 0 - 1 T. atacamensis 1 0 - 1 6 1 1 - 0 15 1 0 - 1 19 73 0.250 1 0 - 1 4 1 0 -+ 1 Tapinurus semitaeniatus 1 0 - 1 T. melanopleurus 1 0 - 1 Uracentron azureum 74 1.000 1 0 - 1 4 75 1.000 1 0 - 1 18 1 1 - 2 19 76 0.500 1 0 - 1 Uranoscodon superciliosus 1 0 - 1 Plica umbra 77 1.000 1 0 - 1 Strobilurus torquatus 19921992FROST: TROPIDURUS GROUP OF LIZARDS 65

APPENDIX 4 Apomorphy List by Stem (Taxon) Noted in Figure 33 Transformations noted by a double dagger (t) are of equally parsimonious placement elsewhere. In the case of terminal taxa even though the character assignment is not equivocal, the plesiomorphic member of the transformation pair may have alternative placement in the topology of the tree. Branch Trans. series Steps ci Change 1 11 1 1.000 0-. 1 12 1 1.000 0-. 1 27 1 1.000 28 0.250 44 1.000 62 0.333 0 1 67 1 1.000 0 - 1 Uranoscodon supercilosus 2 1 0.500 4 1 0.500 0- 1 5 1 0.500 1 19 1 0.250 0- 22 1 0.250 23 1 0.500 30 1 0.333 1 1.000 37 O-b 1 45 1 0 - 0.500 0-. 1it 46 1 0.333 47 1 0.333 48 1 0.571 0 - 51 1 0.250 I 57 2 0.250 0 - 1 59 1 0.333 0 4I 64 1 0.500 0-. 1 0 - 2 70 1 0.500 0-b 1 71 0.500 0 - 3 0-. 1 76 0.500 0 1 0-. 1 2 13 1.000 33 1.000 0 1 3 43 1.000 4 48 0.571 0 1 0-. 1 53 1.000 0 1 O-> 1 60 1.000 O-& 1 71 1 0.500 0 1 73 1 0.250 74 1 1.000 0-.1 - 21 T. atacamensis 72 0.250 0-. 1 T. stolzmanni 47 1 0.333 T. bivittatus 17 0.500 00-.- 1 5 1 1.000 0-. 1 7 1.000 0-b 1 12 1.000 1 - 2 14 1.000 15 0.500 0-.0 1I 16 1.000 17 0.500 0-. 1 20 1.000 0-.- 1 30 0.333 1 2 33 1.000 O - I 34 1.000 O - I 35 1.000 66 AMERICAN MUSEUM NOVITATES NO. 3033

APPENDIX 4 Apomorphy List by Stem (Taxon) Noted in Figure 33-(Continued) Branch Trans. series Steps ci Change 42 1 0.400 0 it 48 1 0.571 0 2 T. nanuzae 22 1 0.250 0 - 1 26 1 1.000 0 1 50 1 1.000 0b 1 6 25 1 0.333 0 - 1 42 1 0.400 1 - 2t 45 1 0.500 0B 1 72 1 0.250 0 1 7 49 1 0.500 0 1 54 1 0.500 0 1 8 56 1 0.333 0 I1 T. itambere 68 1 0.500 0- 1 9 31 1 0.500 0b 1 10 55 1 1.000 0 1 T. hispidus 21 1 0.333 0 I1 11 56 1 0.333 0 1 T. montanus 49 1 0.500 1 2 12 51 1 0.250 0 I1 T. torquatus 49 1 0.500 '1 0 13 29 1 0.667 0b 1 35 1 1.000 Ib 2 41 1 0.667 0 - 1t 48 1 0.571 2 - it 52 1 0.500 0 - 1 63 1 1.000 0 I1 71 1 0.500 0B 2 14 6 1 1.000 0b 1 24 1 0.250 0 1 39 1 1.000 0 - I Tapinurus semitaeniatus 3 1 1.000 0 1 6 1 1.000 I 2 18 1 0.500 0b 1 19 1 0.250 0 1 21 1 0.333 0 1 29 1 0.667 1 - 2 31 1 0.500 1 - 0 32 1 1.000 0 1 73 1 0.250 0 I1 T. bogerti 15 1 0.500 1 - 0 28 1 0.250 1> 0 57 1 0.250 0b 1 62 1 0.333 1¢ 0 15 42 1 0.400 2 1 47 1 0.333 0 1 48 1 0.571 1 3 54 1 0.500 1 - 0 56 1 0.333 1 - 0 59 1 0.333 0 - 1 65 1 1.000 0 1 72 1 0.250 1 0 T. melanopleurus 73 1 0.250 0 I1 1992 FROST: TROPIDURUS GROUP OF LIZARDS 67

APPENDIX 4 Apomorphy List by Stem (Taxon) Noted in Figure 33-(Continued) Branch Trans. series Steps ci Change T. spinulosus 17 1 0.500 2 - 0 28 1 0.250 1- 0 41 1 0.667 1 0:1o 47 1 0.333 1- 2 58 1 0.500 0- 1 16 5 1 0.500 0- 1 8 1 1.000 0- 1 19 1 0.250 0- lt 21 1 0.333 0 - 1 23 1 0.500 0- 1 24 1 0.250 0- 1t 29 1 0.667 1- 0 41 1 0.667 1- 2 57 1 0.250 0- 1t 70 1 0.500 0 - 1 17 10 1 1.000 0- 1 22 1 0.250 0- 1 47 1 0.333 1- 2 58 1 0.500 0- 1 64 1 0.500 0- 1 69 1 1.000 0- 1 Plica plica 24 1 0.250 1 - Ot 46 1 0.333 0 - 1 50 1 1.000 0 - 2 63 1 1.000 1 - 2 P. umbra 2 1 0.500 0 - 1 4 1 0.500 0- 1 19 1 0.250 1 - °t 25 1 0.333 1- 0 42 1 0.400 1- 0 51 1 0.250 1- 0 52 1 0.500 1- 0 70 1 0.500 1- 2 71 1 0.500 2- 3 76 1 0.500 0- 1 18 38 1 1.000 0- 1 48 1 0.571 3- O: 75 1 1.000 0- 1 S. torquatus 47 1 0.333 1- 0 57 1 0.250 1 o0t 77 1 1.000 0- 1 19 17 1 0.500 2t 0 18 1 0.500 0- 1 25 1 0.333 1- 0 30 1 0.333 1b 0 36 1 1.000 0- 1 40 1 1.000 0 - 1 42 1 0.400 1 - 0 46 1 0.333 0- 1 48 1 0.571 0- it 59 1 0.333 1- 0 61 1 1.000 0- 1 68 AMERICAN MUSEUM NOVITATES NO. 3033

APPENDIX 4 Apomorphy List by Stem (Taxon) Noted in Figure 33-(Continued) Branch Trans. series Steps ci Change 62 1 0.333 1¢ 0 66 1 1.000 0b 1 68 1 0.500 0b 1 71 1 0.500 2. 3 72 1 0.250 0 1 75 1 1.000 Ib 2 Uracentron azureum 9 1 1.000 0 I 28 1 0.250 1 0 51 1 0.250 1 0 73 1 0.250 0 I Uracentronflaviceps 22 1 0.250 0 a 1 24 1 0.250 I of T. insulanus 71 1 0.500 0 I1 T. oreadicus 49 1 0.500 1 - 2

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